tag:blogger.com,1999:blog-32484128038147302502016-12-04T06:15:29.839-08:00The Theropod Database BlogHere's a place where I can post my thoughts on new papers, provide updates on my projects, and post info that will eventually be on my website The Theropod Database - <a href="http://theropoddatabase.com/">http://theropoddatabase.com/</a> . It will center on theropods, but may delve into other topics as well such as phylogenetics.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.comBlogger220125tag:blogger.com,1999:blog-3248412803814730250.post-20280661357859217742016-08-31T03:52:00.002-07:002016-08-31T03:52:59.021-07:00The Monster of Minden published at lastThose of you who frequented the Dinosaur Mailing List in the early 2000s might remember references to a supposedly gigantic German theropod- Das Monster von Minden.&nbsp; After a few news stories, it was quickly forgotten until an abstract published last year (Rauhut et al., 2015) confirmed it as a new megalosaurid.&nbsp; As of today it has been officially published and described as <i>Wiehenvenator albati</i> (Rauhut et al., 2016).&nbsp;<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://1.bp.blogspot.com/-tF3fik1i-qc/V8auQ4OGYAI/AAAAAAAAA3Q/YM_BISJ4C8w8oO6QHM3q2riWsbWypMi9ACLcB/s1600/Wiehenvenator%2Bskull.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="157" src="https://1.bp.blogspot.com/-tF3fik1i-qc/V8auQ4OGYAI/AAAAAAAAA3Q/YM_BISJ4C8w8oO6QHM3q2riWsbWypMi9ACLcB/s320/Wiehenvenator%2Bskull.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Skull of <i>Wiehenvenator albati</i> holotype (WMN P27275, P27504, P27457, P27477, P27470, P27461, P27462 and P27466), scale equals 100 mm (after Rauhut et al., 2016).</td></tr></tbody></table>The description is generally excellent, with multiple colored high resolution views of each element.&nbsp; Anatomically, <i>Weihenvenator </i>is basically <i>Torvosaurus</i>, so is not that intriguing.&nbsp; Rauhut et al. are the first I know to alter Carrano et al.'s basal tetanurine analysis however- "Several character definitions of the original list of Carrano et al. (2012), especially of cranial characters, were revised and taxa recoded correspondingly." "A complete documentation of the changes in character definition and character codings will be presented elsewhere (Rauhut and Pol, in prep.)."&nbsp; So that's interesting.&nbsp; They also "added the basal tyrannosaur <i>Guanlong </i>to the matrix in order to improve taxon and character sampling in coelurosaurs", which is good but not nearly enough to fix that issue.&nbsp; They also deleted "the Chinese theropod <i>Leshansaurus</i>, as neither Carrano et al. (2012) nor we studied this taxon personally, and the published description (Li et al., 2009) is in Chinese, so codings could only be based on the sparse illustrations."&nbsp; The illustrations are actually rather good, and the Chinese can be translated via Google Translate, so I'd argue against this.&nbsp; In any case, their resulting tree matches Carrano et al.'s except <i>Cryolophosaurus </i>and <i>Sinosaurus </i>are outside Neotheropoda (their Averostra), <i>Chuandongocoelurus </i>is in a trichotomy with ceratosaurs and tetanurines, and <i>Monolophosaurus </i>is closer to megalosaurians than piatnitzkysaurids, all in Megalosauroidea.&nbsp; Several taxa later added to the Carrano et al. matrix are not analyzed (e.g. <i>Chilesaurus</i>, <i>Sciurumimus</i>), which I think would be important when considering basal tetanurine phylogeny.&nbsp; Rauhut was an author of <i>Sciurumimus </i>after all.&nbsp; Finally, their informal supertree in figure 26 list "Anchiornithosaurs" in quote marks as a group sister to Avialae, which is... interesting.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://1.bp.blogspot.com/-KBMk8FLrjYA/V8a1jaRbRPI/AAAAAAAAA3s/ESdkqxRZLf8VL1DXQWmdMfVU-48MMDDdgCLcB/s1600/Rauhut%2Bet%2Bal%2B2016%2Btetanurine%2Bphylo.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://1.bp.blogspot.com/-KBMk8FLrjYA/V8a1jaRbRPI/AAAAAAAAA3s/ESdkqxRZLf8VL1DXQWmdMfVU-48MMDDdgCLcB/s320/Rauhut%2Bet%2Bal%2B2016%2Btetanurine%2Bphylo.jpg" width="192" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Phylogeny found by Rauhut et al. (2016) after a posteriori exclusion of <i>Streptospondylus </i>using a modified version of Carrano et al.'s basal tetanurine matrix (modified after Rauhut et al., 2016).</td></tr></tbody></table>But what about its size?&nbsp; That was the big thing about the Minden Monster back in the day, with some paleontologists estimating its length as high as ~15 m (<a href="http://www.miketaylor.org.uk/dino/faq/s-size/predator/index.html#9">http://www.miketaylor.org.uk/dino/faq/s-size/predator/index.html#9</a>).&nbsp; Back in 2001 I estimated its size as ~7-8 m (<a href="http://dml.cmnh.org/2003Jul/msg00355.html">http://dml.cmnh.org/2003Jul/msg00355.html</a>) based on maxillary and fibular dimensions.&nbsp; Rauhut et al. don't do a convincing job of calculating this.&nbsp; They say "the maxilla is c. 82% of the size of that of <i>Torvosaurus</i> <i>gurneyi</i>, which was estimated to be approximately the size of <i>Gorgosaurus </i>or <i>Daspletosaurus </i>(c. 10 m in length and 4 to 5 tons in weight) by Hendrickx and Mateus (2014a)."&nbsp; So that'd be ~8 meters long.&nbsp; They then say "On the other hand, the caudal vertebrae are closely comparable in size to elements from a similar position in <i>Torvosaurus tanneri</i>, and the fibulae are even slightly longer than those referred to the latter taxon (Britt, 1991); this taxon was estimated to be approximately 9 m in body length by Britt (1991). Thus, <i>Wiehenvenator </i>is one of the largest theropods found so far in Europe and might only have been slightly smaller than <i>Torvosaurus gurneyi</i>."&nbsp; Except that there are multiple <i>Torvosaurus</i> individuals present in the collection described by Britt, who never says what his ~9 meter estimate is based on.&nbsp; If you use the scale bar on their skeletal reconstruction, <i>Wiehenvenator </i>is ~8.3 m long.&nbsp; So good job me thirteen years ago.<br /><br /><b>References</b>- <span class="reference-text">Rauhut, </span><span class="reference-text">Hübner and Lanser, 2015. A new theropod dinosaur from the late Middle Jurassic of Germany and theropod faunal turnover during the Jurassic. Libro de resúmenes del V Congreso Latinoamericano de Paleontología de Vertebrados. 62.</span><br /><br />Rauhut, Hübner and Lanser, 2016. A new megalosaurid theropod dinosaur from the late Middle Jurassic (Callovian) of north-western Germany: Implications for theropod evolution and faunal turnover in the Jurassic. Palaeontologia Electronica. 19.2.26A, 1-65.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-16662239846172027592016-07-13T18:10:00.000-07:002016-07-15T23:02:04.278-07:00Is Gualicho Aoniraptor?A very cool theropod was published today- <i>Gualicho shinyae </i>(Apesteguia et al., 2016).&nbsp; It's a large taxon from the Huincul Formation of Argentina, notable for having tiny, didactyl arms and generally resembling <i>Deltadromeus</i>.&nbsp; The authors propose both are related to megaraptorans instead of ceratosaurs... and wait, didn't we just have this situation (Motta et al., 2016)?&nbsp; I might as well be the first person* to propose this seemingly obvious synonymy- <i>Gualicho </i>is the same taxon as the recently named <i>Aoniraptor</i>.&nbsp; Both are from the same formation, of similar size (mid caudals ~87 vs. 83 mm), and <i>Gualicho </i>has two diagnostic characters noted for <i>Aoniraptor</i>- "anterior mid-caudal vertebrae with fan-shaped prezygapophyses lacking a discernible articular surface for contacting the postzygapophyses; presence of a blunt and thick process on the lateral surface of the prezygapophyses of anterior midcaudal vertebrae", and might have the third- "mid-posterior caudals with a pair of nonarticular flat surfaces located on the posterodorsal corner of the centrum."&nbsp; <strike>As <i>Aoniraptor </i>was named first (June 16th vs. July 13th), it has precedent over <i>Gualicho</i>.</strike>&nbsp; <b>Edit #2: As Brad McFeeters notes in a comment below, the volume <i>Aoniraptor </i>was named in has yet to be physically published and does not contain a ZooBank registration, thus <i>Gualicho </i>has precedent.</b><br /><br /><b>*Edit: After posting this, I learned both Andrea Cau (<a href="http://theropoda.blogspot.com/2016/07/nuovi-resti-di-aoniraptor-ehm-benvenuto.html" target="_blank">here</a>) and Brad McFeeters (on Facebook) independently came to the same conclusion.&nbsp; Guess it really was an obvious synonymy. ;)</b><br /><br />I'm skeptical that these and <i>Deltadromeus </i>belong in Tetanurae, as it only takes 4 steps to move them to Ceratosauria in Apesteguia et al.'s version of Carrano's tetanurine analysis, and <i>Deltadromeus </i>wasn't added to their version of Novas' tetanurine analysis which also lacks any ceratosaurs except the outgroup <i>Ceratosaurus</i>.&nbsp; They note "<i>Gualicho </i>does exhibit some apparent ceratosaurian synapomorphies, most notably a robust MT III with posteriorly expanded proximal articulation and a flange-like m. iliofibularis tubercle."&nbsp; It would have been interesting to run them in a ceratosaur analysis.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://1.bp.blogspot.com/-ffwucni20I4/V4bfb-rAG9I/AAAAAAAAA20/sEYl-6jD4WADnmyrBNfd21Cmb0cmGBLdgCLcB/s1600/Aoniraptor%2Bvs%2BGualicho.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="163" src="https://1.bp.blogspot.com/-ffwucni20I4/V4bfb-rAG9I/AAAAAAAAA20/sEYl-6jD4WADnmyrBNfd21Cmb0cmGBLdgCLcB/s320/Aoniraptor%2Bvs%2BGualicho.jpg" width="320" />&nbsp;</a></td><td style="text-align: center;"></td><td style="text-align: center;"></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotypes of <i>Gualicho shinyae</i> (MPCN PV 0001; top) and <i>Aoniraptor libertatum</i> (MPCA-Pv 804; bottom) in posterior (E), anterior (F), left lateral (G) and dorsal (H) views.&nbsp; Modified from Apesteguia et al. (2016) and Motta et al. (2016).</td><td class="tr-caption" style="text-align: center;"><br /></td><td class="tr-caption" style="text-align: center;"><br /></td></tr></tbody></table>&nbsp;Wonder where they go in the Lori analysis....<br /><br /><b>References</b>- Apesteguía, Smith, Juárez Valieri and Makovicky, 2016. An unusual new theropod with a didactyl manus from the Upper Cretaceous of Patagonia, Argentina. PLoS ONE. 11(7), e0157793.<br /><br />Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson Egli and Novas, 2016. <span class="reference-text">New theropod fauna from the Upper Cretaceous (Huincul Formtation) of northwestern Patagonia, Argentina. In Khosla and Lucas (eds.). Cretaceous period: Biotic diversity and biogeography. New Mexico Museum of Natural History and Science Bulletin. 71, 231-253.</span><br /><br /><br />Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com18tag:blogger.com,1999:blog-3248412803814730250.post-642437700866706402016-07-06T18:09:00.001-07:002016-07-07T12:30:35.044-07:00Is Lagerpeton a proterochampsian? The Dromomeron angleSorry for the lack of posts (or Database updates) lately, as I've been working with Scott Hartman to add Lori to the coelurosaur matrix and finalizing that study.&nbsp; But I started writing this as a DML reply and figured why not make it a post instead...<br /><br />Novas and Agnolin (2015) have an abstract on <i>Lagerpeton </i>that proposes it is actually a proterochampsian sister to <i>Tropidosuchus</i>.&nbsp; If true, this would be a rare time David Peters got something right, as he had this idea back in 2011- <a href="https://pterosaurheresies.wordpress.com/2011/10/15/what-is-lagerpeton-the-heretical-view/">https://pterosaurheresies.wordpress.com/2011/10/15/what-is-lagerpeton-the-heretical-view/</a> .&nbsp; I hope Novas and Agnolin credit him if they write a paper on this.<br /><br />However, something not addressed by Novas and Agnolin is how <i>Dromomeron </i>relates.&nbsp; According to Nesbitt (2011), <i>Tropidosuchus </i>lacks all lagerpetonid characters shared by <i>Lagerpeton </i>and <i>Dromomeron</i>- Anterolateral tuber of the proximal portion of the femur absent, the anterolateral face is flat (302-1), Femoral head hook shaped in medial and lateral views (306-1), Ventral emargination present on anterolateral side of the femoral head (310-1), Crista tibiofibularis larger than the medial condyle (326-1), Dorsally expanded process on the posterolateral portion of the tibial facet of the astragalus expanded into a distinct, raised process (5 posterior ascending process of Sereno and Arcucci, 1994a) (355-1), Concave articular surface for the fibula on the calcaneum (378-2).&nbsp; Of the proterochampsian characters noted by Novas and Agnolin as present in <i>Lagerpeton</i>, <i>Dromomeron </i>lacks two- femoral 4th trochanter proximodistally expanded; caudal surface of distal tibia with a middle tubercle surrounded by two shallow concavities; but has one- astragalus with anteromedial corner acute.&nbsp; Others are unknown since <i>Dromomeron </i>is so fragmentary, but at least "elongate and compact metatarsus with metatarsal V reduced and devoid of phalanges" would be expected in a taxon close to <i>Marasuchus</i> (which has the conditions), so that works for either placement of <i>Lagerpeton</i>.&nbsp; Then there are the dinosauriform-like characters of <i>Lagerpeton </i>scored by Nesbitt as absent in <i>Tropidosuchus</i>- Straight cnemial crest (328-1); Longest metatarsal longer than 50% of tibial length (383-1); Metatarsal V "hooked" proximal end absent, and articular face for distal tarsal 4 subparallel to shaft axis (398- 1).&nbsp; And the archosaurian characters of <i>Lagerpeton </i>scored as absent in <i>Tropidosuchus</i>- Articular surfaces for fibula and distal tarsal IV on the calcaneum continuous (380-1), Anteromedial tuber of the proximal portion of the femur present (300-1), Tibial facet of the astragalus divided into posteromedial and anterolateral basins (366-1).<br /><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://4.bp.blogspot.com/-E4pkdfmcf1g/V32nwzyxOsI/AAAAAAAAA2c/RenSXHEztPkLP8td_pEuh8YIdcKZK6cBACLcB/s1600/Lagerpeton%2Bmove.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://4.bp.blogspot.com/-E4pkdfmcf1g/V32nwzyxOsI/AAAAAAAAA2c/RenSXHEztPkLP8td_pEuh8YIdcKZK6cBACLcB/s320/Lagerpeton%2Bmove.jpg" width="298" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Phylogeny of eucrocopodans recovered by Ezcurra (2016), with Novas and Agnolin's (2015) proposed reidentification of <i>Lagerpeton </i>indicated in red (modified from Ezcurra, 2016).</td></tr></tbody></table><br />Taking all of this into account (with the caveats that I haven't checked the accuracy of any characters except the proterochampsian states of <i>Dromomeron</i>, and that this only uses unambiguous characters listed by Nesbitt), Novas and Agnolin have ten characters that might need to converge if <i>Lagerpeton </i>is a dinosauriform relative, but there are nine characters that might need to converge if <i>Lagerpeton </i>is a proterochampsian (the seven from Nesbitt listed here plus two listed by Novas and Agnolin).&nbsp; So that's one step in favor of a proterochampsian identity.&nbsp; But if <i>Dromomeron </i>is a proterochampsian too, that adds three more steps (the two listed above plus dinosauromorph character 313 of Nesbitt that is present in <i>Dromomeron </i>but unknown in <i>Lagerpeton</i>).&nbsp; Thus dinosauriform-relative <i>Lagerpeton </i>is two steps shorter.&nbsp; And if <i>Dromomeron </i>is a dinosauriform relative but <i>Lagerpeton </i>is not, that adds five steps (the six lagerpetonid characters that would need to be convergent minus the proterochampsian-like acute astragalar corner).<br /><br />So basically, ignoring <i>Dromomeron </i>might make proterochampsian <i>Lagerpeton </i>a slightly better option but a proterochampsian <i>Dromomeron </i>turns those odds around, and a proterochampsian <i>Lagerpeton </i>but a dinosauriform-relative <i>Dromomeron </i>is even worse.&nbsp; So pending further analysis, I favor the traditional view (note Ezcurra's huge 2016 archosauromorph analysis which sampled proterochampsian diversity also recovered this topology).&nbsp; If <i>Lagerpeton </i>is a proterochampsian, then Lagerpetonidae is a junior synonym of Proterochampsia definition-wise (and Proterochampsidae ICZN-wise).&nbsp; However, Novas and Agnolin are wrong in claiming "The exclusion of <i>Lagerpeton </i>from the dinosaur lineage results in the removal of the clade Dinosauromorpha, which was originally conceived to encompass <i>Lagerpeton </i>plus Dinosauriformes."&nbsp; The only proposed definition of Dinosauromorpha that mentions <i>Lagerpeton </i>is one of Sereno's (1991) which is "<i><i>Lagerpeton</i> chanarensis</i>, <i><i><i>Lagosuchus</i> talampayensis</i></i>, <i><i>Pseudolagosuchus</i> major</i>, Dinosauria (inc. Aves), and all descendants of their common ancestor."&nbsp; Following this definition, Dinosauromorpha merely moves stem-ward to be the Proterochampsia+Archosauria clade which is AFAIK otherwise unnamed.&nbsp; The other proposed definitions are all basically "taxa closer to dinosaurs than pterosaurs", so Dinosauromorpha would stay where it is even if <i>Lagerpeton </i>isn't a member.<br /><br /><b>References</b>- Sereno, 1991. Basal archosaurs: Phylogenetic relationships and functional implications. Society of Vertebrate Paleontology Memoir. 2, 1 53 pp<br /><br />Nesbitt, 2011. The early evolution of archosaurs: Relationships and the origin of major clades. Bulletin of the American Museum of Natural History. 352, 292 pp. <br /><br />Novas and Agnolin, 2015. <i>Lagerpeton chanarensis</i> Romer (Archosauriformes): A derived proterochampsian from the Middle Triassic of NW Argentina. Libro de resúmenes del V Congreso Latinoamericano de Paleontología de Vertebrados. 48.<br /><br />Ezcurra, 2016. The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms. PeerJ. 4:e1778.<br /><i><br /></i>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com1tag:blogger.com,1999:blog-3248412803814730250.post-10824212119986163552016-04-02T02:31:00.001-07:002016-04-23T11:41:18.644-07:00Database update plus Predatory Dinosaurs of the World with phylogenetic nomenclatureHey fellow 80s kids!&nbsp; Like me, you probably grew up with Paul's (1988) Predatory Dinosaurs of the World as your first serious non-technical theropod source, along with The Dinosauria.&nbsp; After <a href="http://theropoddatabase.blogspot.com/2016/03/peters-theropod-craziness.html" target="_blank">raking Peters' horrific theropod phylogeny over the coals and applying current phylogenetic nomenclature for fun</a>, I wondered how the most famous non-technical topology of our time would fare.&nbsp; So here's Paul's phylogeny from his influential book using the taxonomy from The Theropod Database.&nbsp; Note a number of his named groups are paraphyletic in his topology (based on overleaf 10-1, with additional details based on his comments in the taxonomic section).&nbsp; Only species with entries were included.<br /><span style="font-size: x-small;"><br /></span><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">Dinosauromorpha <br />Dinosauriformes</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">|*-Saltopus<br />|--Lagerpetonidae<br />|&nbsp; |--Lagosuchus (inc. Marasuchus)<br />|&nbsp; |--Lewisuchus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">|&nbsp; `--Lagerpeton<br />`--Herrerasauridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp; |--Staurikosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp; `--+--Ischisaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Herrerasauria</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |?*Alwalkeria</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |?-Protoavis</span></span> </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Freguellisaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--Aliwalia</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Herrerasaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Avepoda</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Procompsognathus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Megalosauria</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Spinosauroidea (= Coelophysoidea, Spinosauridae)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |?-Elaphrosaurus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Coelophysidae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; Coelophysis</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; |--rhodesiensis</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; `--bauri</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Dilophosauridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |--Liliensternus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--+--Dilophosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Baryonychinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Baryonyx</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Spinosaurinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; Spinosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Neotheropoda (= Averostra, Neoceratosauria, Abelisauroidea, Torvosauridae)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Ceratosauria (= Ceratosauridae)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Sarcosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Ceratosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; |?-ingens</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; `--nasicornis </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Tetanurae (= Abelisauridae, Orionides)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Megalosauroidea (= Megalosauridae) </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--+--Megalosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; |--Torvosaurus&nbsp;</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; `--Poekilopleuron</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Abelisauria</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |?-Noasauridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Noasaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |--Carnotaurinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Carnotaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Abelisaurinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; Abelisaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Eustreptospondylinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Eustreptospondylus (inc. Streptospondylus)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Piatnitzkysauridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Piatnitzkysaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Gasosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Marshosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Allosauroidea</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Metriacanthosauridae (= Sinraptoridae)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; |--Yangchuanosaurus (inc. Metriacanthosaurus? sp.) </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Metriacanthosaurus (assuming Sinraptor)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Avetheropoda (= Neotetanurae, Allosauria, Tyrannoraptora)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |*-Dryptosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Compsognathidae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Compsognathus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Tyrannosauroidea (= Carnosauria)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |--Coeluridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; |--Aristosuchus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; `--Coelurus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |--Proceratosauridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; |--+--Proceratosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; |&nbsp; `--Pivetaeusaurus&nbsp;</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; |--Rapator</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; `--Ornitholestes</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--+--Allosauridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Allosaurus fragilis (inc. A. atrox, A. amplexus)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; `--Carcharodontosauridae (= Carcharodontosauria)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--+--Chilantaisaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--Shaochilong </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--+--+--Acrocanthosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; `--Becklespinax </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--Indosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--Labocania</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--Erectopus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--Carcharodontosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--Bahariasaurus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--Aublysodontinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Aublysodon</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |--Shanshanosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--Stygivenator </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--Indosuchus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--Alioramini</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Alioramus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--Tyrannosauridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Albertosaurinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; |--Alectrosaurus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+*-Albertosaurus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |--Nanotyrannus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--+--Gorgosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--+--arctunguis</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--Dinotyrannus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--Tyrannosaurinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Daspletosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Tarbosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Tyrannosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Coelurosauria (= Aviremigia, Ornithes)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Archaeopterygidae (= Saurornithes)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Archaeopteryx</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Eudromaeosauria</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Dromaeosauridae (= Dromaeosaurinae)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Dromaeosaurus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `?-Adasaurus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Eumaniraptora (= Pennaraptora, Averaptora)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Deinonychosauria (= Velociraptorinae)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Deinonychus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--Saurornitholestes</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Velociraptor</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Avialae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |?-Ornithomimosauria (= Arctometatarsalia)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Harpymimus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Ornithomiminae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |?-Deinocheiridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Deinocheirus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |--Garudimimus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Ornithomimidae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |*-affinis</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |*-Archaeornithomimus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--+--Struthiomimus&nbsp;</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |?-Ornithomimus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Dromiceiomimus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--brevitertius</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--samueli</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Gallimimus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Oviraptorosauria (= Caenagnathiformes)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Caenagnathoidea</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Caenagnathidae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; |--Microvenator </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; `--+--Caenagnathinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Chirostenotes</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; | &nbsp; &nbsp; `--Elmisaurinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; Elmisaurus </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Oviraptoridae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |--Oviraptorinae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Oviraptor (assuming Citipati)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--"Ingeniinae"</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; "Ingenia"</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Maniraptora (= Avimimidae, Paraves)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Troodontidae</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Troodon</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; |--Saurorni<span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">thoides</span></span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; |--Hept<span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">asteornis</span></span></span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; `?-Brad<span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">ycneme</span> </span>&nbsp;</span> </span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Avimimus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Ornithothoraces (= Avebrevicauda, Ornithopectae)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Enantiornithes</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Iberomesornis (assuming Enantiornis)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Ichthyornithes (= Ambiortiformes)</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Ambiortus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Ichthyornis</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Gansuiformes</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Gansus</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Hesperornithes</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Hesperornis</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Aves</span></span><br /><span style="font-size: x-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"><br /></span></span>So how does it fare?&nbsp; Pretty well, though Herrerasauridae, Megalosauria, Spinosauridae and Tyrannosauridae have definitions that fail.&nbsp; There are some uncertainties based on the unclear position of ornithomimosaurs, and due to the date there are no taxa between <i>Archaeopteryx </i>and ornithothoracines.&nbsp; But overall, not bad.&nbsp; <br /><br />Among the "Theropod Odds and Ends", Taquet's (1984) taxon has yet to be redescribed but seems to be a coelophysoid, <i>"Labrosaurus" ferox</i> is agreed to be an allosaurid/<i>Allosaurus </i>specimen and maybe a portion of the <i>A. fragilis</i> neotype.&nbsp; (<i>Magnosaurus</i>) <i>nethercombensis </i>is a megalosauroid, possibly afrovenatorine.&nbsp; (<i>Duriavenator</i>) <i>hesperis </i>is a megalosaurid.&nbsp; <i>Xuanhanosaurus </i>seems to be a metriacanthosaurid.&nbsp; <i>Iliosuchus </i>may be a basal tyrannosauroid or other basal coelurosaurian, which is debated for taxa known from far more than only ilia.&nbsp; <i>Stokesosaurus </i>is universally agreed to be a basal tyrannosauroid.&nbsp; Finally, <i>Segisaurus </i>is agreed to be a coelophysoid closer to <i>Coelophysis </i>than <i>Dilophosaurus</i>.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://3.bp.blogspot.com/-D5zacDU5qFU/Vv-MJ5izm_I/AAAAAAAAA2E/xuuYzV54KCE81nRHU_nePG8ERaxu-gfhQ/s1600/Segnosaurus%2Btricarinate.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="229" src="https://3.bp.blogspot.com/-D5zacDU5qFU/Vv-MJ5izm_I/AAAAAAAAA2E/xuuYzV54KCE81nRHU_nePG8ERaxu-gfhQ/s320/Segnosaurus%2Btricarinate.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Twenty-third (II) and twenty-second (III) dentary tooth of holotype of <i>Segnosaurus galbinensis</i> (IGM 10/80), after Zanno et al. (2016). Note the weird lingual carina (lc).</td></tr></tbody></table>Oh, and I updated the Database this month, but less than usual due to coding more in the Lori matrix.&nbsp; Besides the last sauropodomorph entry&nbsp; (<i>Ignavusaurus </i>?= <i>Massospondylus</i>), a couple braincases were added to the Bissekty tyrannosauroid(s?) and named <i>Timurlengia</i>.&nbsp; Wonder where this falls out in the new tyrannosauroid matrix of Brusatte and Carr (2016).&nbsp; The Pierre Shale hesperornithoids of Manitoba were clarified by Aotsuka and Sato (2016), with the description of a new species.&nbsp; <i>Segnosaurus</i>' mandible was also redescribed (Zanno et al., 2016), which I hope is the first of many therizinosaurian redescriptions from them.&nbsp; It's quite an odd specimen, exhibiting a coronoid process like oviraptorosaurs, no posterior surangular ridge or foramen and TRIcarinate posterior dentary teeth.&nbsp; Yes, unlike (?)any other theropod, <i>Segnosaurus </i>has a lingual carina with serrations (see above).<br /><br /><b>References</b>- Taquet, 1984. Two new Jurassic specimens of coelurosaurs (Dinosauria). In Hecht, Ostrom, Viohl and Wellnhofer (eds.). The Beginnings of Birds: Proceedings of the International Archaeopteryx Conference, Eichstaett. 229-232.<br /><br />Paul, 1988. Predatory Dinosaurs of the World. Simon &amp; Schuster. 464 pp. <br /><br />Aotsuka and Sato, 2016. Hesperornithiformes (Aves: Ornithurae) from the Upper Cretaceous Pierre Shale, southern Manitoba, Canada. Cretaceous Research. doi: 10.1016/j.cretres.2016.03.003.<br /><br />Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports. 6, 20252.<br /><br />Zanno, Tsogtbaatar, Chinzorig and Gates, 2016. Specializations of the mandibular anatomy and dentition of <i>Segnosaurus galbinensis</i> (Theropoda: Therizinosauria). PeerJ. 4:e1885.<br /><br />Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com1tag:blogger.com,1999:blog-3248412803814730250.post-88062560023179745632016-03-26T01:43:00.004-07:002016-03-26T01:43:54.238-07:00Peters' theropod crazinessSo David Peters, whose analyses I have <a href="http://theropoddatabase.blogspot.com/2013/02/testing-peters-dinosaur-phylogeny.html" target="_blank">thoroughly trashed</a> <a href="http://theropoddatabase.blogspot.com/2013/03/peters-dinosaur-phylogeny-fails-finale_29.html" target="_blank">on this blog before</a>, has branched out into theropods.&nbsp; In my prior foray into his <a href="http://theropoddatabase.blogspot.com/2012/06/why-doesnt-peters-find-dinosauria-or.html" target="_blank">basal dinosaurian</a> <a href="http://theropoddatabase.blogspot.com/2013/04/are-daemonosaurus-silesaurids-and.html" target="_blank">phylogeny</a>, I was surprised by how poorly coded (~30% miscoded) taxa were for his tiny and badly formed analysis.&nbsp; Now that he's trying to interpret MY turf, it's just sad.&nbsp; Or laughable.&nbsp; Or laughably sad, take your pick.&nbsp; Even in the best of circumstances, taking 228 characters designed for amniote phylogeny and coding theropods for them is likely to give you a terrible result.&nbsp; But add Peters' DGS to the mix, where he takes photos and discovers missing bones in a slab or traces his own outlines, and you have hilarious fiction.&nbsp; Peters might know pterosaurs, but when he tries tracing theropods, it's obvious he does NOT know theropods.&nbsp; He mixes bone outlines, identifies random sediment texture or feathers as bones, and based on his results just has no idea how theropods look.&nbsp; I commented on this on his <i>Archaeornithura </i>skull post last week (see last figure below), but Peters deleted the comment and changed his reconstruction to be wrong in different ways (new ways include that the pubis is the ischium, the ischium has a huge obturator process, the manus has a complete digit, as does the fifth pedal digit(!)).<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://1.bp.blogspot.com/-eblftydTRtU/VvY_JoPK25I/AAAAAAAAA1Y/YWhOdKdKFLQcw_zqXGNNPxYr1e6l3RH9Q/s1600/Sinosauropteryx%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="202" src="https://1.bp.blogspot.com/-eblftydTRtU/VvY_JoPK25I/AAAAAAAAA1Y/YWhOdKdKFLQcw_zqXGNNPxYr1e6l3RH9Q/s320/Sinosauropteryx%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Reconstructions of <i>Sinosauropteryx prima</i> by me (top), and Peters (bottom), scaled to same femoral length.</td></tr></tbody></table>As an example, here's <a href="http://www.reptileevolution.com/sinosauropteryx.htm" target="_blank"><i>Sinosauropteryx</i> as reconstructed by him</a> vs. by me.&nbsp; Now, Peters is no doubt the more experienced artist, and my reconstruction isn't perfect.&nbsp; I made it back in 2005, and if I were to redraw it today, I'd give the tibia and fibula less generic shapes (e.g. larger cnemial crest, flatter distal tibia, expanded proximal fibula), orient the humerus to not be in posterior view, and get rid of the post-obturator notch in the ischium, which was later found to be erroneous.&nbsp; But compare that to Peters' monstrosity.&nbsp; The squamosal is just a triangular plate, lacking any posterior process so that the quadrate head is impossibly just floating freely posterior to it.&nbsp; The quadrate has a narrow orbital/pterygoid process unlike anything but derived birds.&nbsp; There are twenty-five presacral vertebrae unlike (?)all non-maniraptoran avepods, the caudals vary randomly in length by large degrees unlike any theropod, and the chevrons are all short as if it were a derived paravian.&nbsp; In the pectoral girdle, there's some huge crescent where the sternum would be, but <i>Sinosauropteryx </i>lacks an ossified sternum.&nbsp; The ilium is just wacky- tiny postacetabular process and concave dorsal margin.&nbsp; The ischium is a thick blob with nary an obturator process.&nbsp; Perhaps most sadly, Peters can't even draw the tibia as longer than the femur, which is (?)universal in small theropods, stated even in the crappy original Chinese description and is character 195 in Peters' analysis.<br /><br />With his reconstructions being more or less fictional animals, it's no surprise the cladograms based on them will be equally unrealistic.&nbsp; And lo, they are!&nbsp; For fun, I give <a href="https://pterosaurheresies.wordpress.com/2016/03/14/the-origin-of-dinosaurs-x2-2010-revisited/" target="_blank">his cladogram from March 14th</a> and applied phylogenetic nomenclature to see how it fares.&nbsp; I listed the assumed position of taxa not yet included, as they are specifiers of various clades.&nbsp; I also list where family-level taxa would have to be renamed due to ICZN rules.<br /><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">Theropoda</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">|--Tawa</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">`--+--+--Staurikosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp; |&nbsp; `--+--Segisaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp; | &nbsp; &nbsp; `--+--Guaibasaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--+--Marasuchus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--Procompsognathus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp; `--Avepoda</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; |--Coelophysoidea</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; |&nbsp; |--Dracoraptor</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; |&nbsp; `--+--Coelophysidae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; |&nbsp; Coelophysis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Dilophosauridae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; Dilophosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; `--Neotheropoda (= Orionides, Avetheropoda, Neotetanurae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |--</span></span><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">Ceratosauria<span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"> (= </span>Spinosauroidea, Carnosauria, Deinocheiridae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Megalosauri<span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">a</span> </span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |--Spinosauridae (= Proceratosauridae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; |--Proceratosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |&nbsp; `--+--Deinocheirus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; | &nbsp; &nbsp; `--+--Xiongguanlong</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--+--Sinocalliopteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--Suchomimus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--Spinosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--+--+--Dilong</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; |&nbsp; `--Guanlong</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; `--Allosauroidea (= Neoceratosauria) (would need to be renamed Megalosauroidea)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |--Abelisauroidea (= Metriacanthosauridae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |--Yutyrannus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--+--Sinraptor (assuming Metriacanthosaurus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; `--Abelisauridae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; Majungasaurus (assuming Abelisaurus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--Ceratosauridae<span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"> </span></span></span><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">(would need to be renamed Megalosauridae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; <span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">Allosauria</span> </span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--Allosauridae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; Allosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Megalosauridae (= Megalosauroidea, Carcharodontosauridae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--+--Sinosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Monolophosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Acrocanthosaurus (assuming Carcharodontosaurus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Eustreptospondylus (assuming Megalosaurus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Ceratosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Coelurosauria (= Tetanurae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; Maniraptoriformes (= Aviremigia, </span><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">Tyrannoraptora, Bullatosauria)</span> </span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Compsognathidae (= Tyrannosauroidea, Arctometatarsalia, Microraptoria)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Ornithomimosauria</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; |--Compsognathus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; `--Struthiomimus (assuming Ornithomimus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--+--Ornitholestes</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--Microraptor</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Sinornithosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--+--Fukuivenator</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Tianyuraptor</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Zhenyuanlong</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Alioramini</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Alioramus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Tyrannosauridae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Gorgosaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Tyrannosauridae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Maniraptora</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; Pennaraptora (= Chuniaoae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Caenagnathiformes (= Oviraptoriformes)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Therizinosauria</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; |--Falcarius</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; `--+--Jianchangosaurus (assuming Therizinosaurus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Rahonavis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Oviraptorosauria</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |--+--Juravenator</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Sinosauropteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--+--Limusaurus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Khaan (assuming Caenagnathus and Oviraptor)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Paraves</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Coeluridae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Tanycolagreus (assuming Coelurus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Eotyrannus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Metornithes (= Eumaniraptora)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Dromaeosauridae (= Alvarezsauroidea, Alvarezsauria, Deinonychosauria)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--+--Haplocheirus</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |&nbsp; `--Shuvuuia (assuming Alvarezsaurus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--Velociraptor (assuming Dromaeosaurus and Deinonychus)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Balaur</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Avialae (= Averaptora)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Sinornithoides (assuming Troodon)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Jinfengopteryginae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; Jinfengopteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Unenlagiinae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Anchiornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--Aurornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Buitreraptor (assuming Unenlagia)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Sinovenator</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--+--Eosinopteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Xiaotingia</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Thermopolis "Archaeopteryx"</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Ornithothoraces (= Ornithes, Ornithopectae)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Archaeopterygidae (= Saurornithes, Enantiornithes)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Archaeornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--+--Archaeopteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Yixian embryo IVPP V14238</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--+--Protopteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Cathayornis (assuming Iberomesornis and Enantiornis)</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Pengornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Sulcavis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Ornithurae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Omnivoropterygiformes</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--"Archaeopteryx" bavarica</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--+--Mei</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp; `--Scansoriopterygidae</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Scansoriopteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Yi</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Epidexipteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Omnivoropteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Jeholornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Jurapteryx</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Pygostylia</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Confuciusornithiformes</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--Wellnhoferia</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; `--Confuciusornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Avebrevicauda</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Chiappeavis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Sapeornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Archaeornithura</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--+--Ichthyornis</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Aves</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Struthio</span></span><br /><span style="font-size: xx-small;"><span style="font-family: &quot;courier new&quot; , &quot;courier&quot; , monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; `--Gallus (assuming Passer)</span></span><br /><br /><br /><span style="font-family: inherit;"><span style="font-family: &quot;arial&quot; , &quot;helvetica&quot; , sans-serif;"><span style="font-size: small;">That's pretty funny.&nbsp; Nomenclature fails in Allosauroidea and Compsognathidae, due to the weird topologies there where taxa not seen as closely related enough to need their neighbors as external specifiers suffer.&nbsp; Ditto for dromaeosaurids vs. alvarezsaurs.&nbsp; Also for archaeopterygids vs. enantiornithines, which no BAD analysis has ever recovered.&nbsp; <br /><br />As for the topology, there's that sister clade to avepods, whose oldest family name would be Procompsognathidae, though none of the members have phylogenetic definitions attached to them.&nbsp; The division in neotheropods is sort of like Rauhut (2003), who had a huge Carnosauria.&nbsp; The most basal megalosaurians are all actually basal tyrannosauroids.&nbsp; Ornithomimids plus tyrannosaurids is a classic clade from Huene to Holtz, and <i>Compsognathus </i>being there reminds me of Olshevsky (1995) having it as a tyrannosaur.&nbsp; Funny how Peters recovered Enigmosauria, though no doubt for completely different reasons than the dinosaur community, given the inclusion of compsognathids, <i>Limusaurus </i>and <i>Rahonavis</i>.&nbsp; Troodontids and unenlagiines being avialans has been more popular recently (e.g. Agnolin and Novas, 2013), and the <i>Archaeopteryx </i>plus Enantiornithes pairing hearkens back to the BANDits' Sauriurae.&nbsp; Similarly, <i>Confuciusornis </i>being closer to Aves than enants reminds me of Kurochkin's (2006) ideas.&nbsp; And those are the only parallels I can make between Peters' non-consensus phylogeny and science.&nbsp; My head's full of non-traditional phylogenetic proposals, but <i>Deinocheirus </i>as a spinosaur, or microraptorans as basal tyrannosaurs while <i>Velociraptor </i>is by alvarezsaurs?&nbsp; That's just nuts.<br /><br />So David, first I'd say you should start using the names <i>Archaeornis</i>, <i>Jurapteryx </i>and <i>Wellnhoferia </i>if you find these taxa away from the <i>Archaeopteryx </i>holotype.&nbsp; But also, I think it would be amusing if you added the following- <i>Megalosaurus</i>, <i>Hexing</i>, <i>Nothronychus</i>/<i>Erlikosaurus</i>, <i>Caudipteryx</i>, <i>Achillobator</i>, <i>Dromaeosaurus </i>and <i>Patagopteryx</i>.</span></span></span><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://4.bp.blogspot.com/-r0k8p_jA-eA/VvZH84SAoNI/AAAAAAAAA1o/UP0A3VoluN8WW8NO1_iVpMy-ghADcArkg/s1600/Archaeornithura%2BPeters.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="252" src="https://4.bp.blogspot.com/-r0k8p_jA-eA/VvZH84SAoNI/AAAAAAAAA1o/UP0A3VoluN8WW8NO1_iVpMy-ghADcArkg/s320/Archaeornithura%2BPeters.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Peters' reconstruction of <i>Archaeornithura meemannae</i> <a href="https://pterosaurheresies.wordpress.com/2016/03/15/is-this-the-missing-skull-of-the-basal-bird-archaeornithura/" target="_blank">from 3-26-16</a>, preserved in case it's deleted like the first was (though Peters to his partial credit does say "Updated March 16, 2016 with new images. The beak, if present, is ephemeral, questionable. Only two scores changed.").</td></tr></tbody></table><br /><span style="font-family: inherit;"><span style="font-size: small;"><b>References</b>- </span></span><span style="font-family: inherit;"><span style="font-size: small;">Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku Saizensen. 9, 92-119 (part 1); 10, 75-99 (part 2). <br />&nbsp;</span></span><br /><span style="font-family: inherit;"><span style="font-size: small;">Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213. </span></span><span style="font-family: inherit;"><span style="font-size: small;">&nbsp;</span></span><br /><br /><span style="font-family: inherit;"><span style="font-size: small;">Kurochkin, 2006. Parallel evolution of theropod dinosaurs and birds. Entomological Review. 86(suppl. 1), S45-S58.</span></span><br /><br />Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, <i>Anchiornis</i> and Scansoriopterygidae. Springer. 96 pp. Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com9tag:blogger.com,1999:blog-3248412803814730250.post-11739771311181506152016-02-24T19:48:00.003-08:002016-02-24T19:48:57.552-08:00Fukuivenator thoughtsSo we have another new theropod, <i>Fukuivenator paradoxus</i> (Azuma et al., 2016).&nbsp; With <i>Fukuiraptor</i>, <i>Fukuititan </i>and <i>Fukuisaurus </i>also in existence, we just need a <i>Fukuipelta</i>, <i>Fukuimimus </i>and <i>Fukuiornis </i>to round out the Kitadani Formation.&nbsp; This was first announced in an anonymous 2009 msn article (now defunct) and Shibata and Azuma's (2010) SVP abstract.&nbsp; While those sources called it a dromaeosaurid, Azuma et al. interestingly say "it in fact represents a bizarre, basally branching maniraptoran theropod with a large number of autapomorphies."&nbsp; Intriguing.&nbsp; Let's check it out...<br /><br />A partial skull is present (though the jugal, palatine, ?ectopterygoid and pterygoid are unillustrated) as well as a dentary fragment.&nbsp; The weirdest thing is the gigantic premaxilla, which is so odd that I would question its identity and propose it's actually the right maxilla, except that it lacks the pneumatic fossa and foramina present on the left maxilla.&nbsp; Contra the authors, the posterolateral process of the premaxilla is broken so (barring the presence of some really well preserved sutures on the maxilla) it cannot be determined if it extends posterior to the external naris like in dromaeosaurids.&nbsp; Similarly, the supposedly large promaxillary fenestra (cited in the diagnosis as well) is actually small.&nbsp; The text claims the lacrimal is T-shaped, but the figure shows the posterodorsal process is unpreserved, and the character is coded unknown in their matrix.&nbsp; Although the authors claim the frontals have dromaeosaurid-like anterolateral notches and sigmoid supratemporal fossa edges, neither is visible in the figured left element.&nbsp; Amusingly, Azuma et al. say one premaxillary tooth "exhibits a "ridge," on which denticles are absent."&nbsp; Ah, you mean a carina? ;)&nbsp; The other premaxillary tooth is indeed interesting in being D-shaped.&nbsp; Surely the fenestra labeled "IX?" in figure 4&nbsp; is the otic fenestra, and the basipterygoid processes are confusingly labeled as laterosphenoids twice in that figure.&nbsp; The supposed medial eustachian foramina are the paired foramina of the basisphenoid recess.&nbsp; Oddly, neither of the eustachian tube characters are coded, but the basisphenoid recess characters are coded correctly.&nbsp; It's an interesting braincase, with no obvious dorsal or posterior tympanic recesses, a reduced basisphenoid recess and laterally diverging basipterygoid processes, most of which are troodontid-y.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://3.bp.blogspot.com/-LSlSHD2hK7g/Vs55fA3LP-I/AAAAAAAAA1E/Ku1Yy4-M6Nk/s1600/Fukuivenator%2Bskull.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="198" src="https://3.bp.blogspot.com/-LSlSHD2hK7g/Vs55fA3LP-I/AAAAAAAAA1E/Ku1Yy4-M6Nk/s320/Fukuivenator%2Bskull.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Reconstructed skull of <i>Fukuivenator paradoxus</i> holotype, premaxilla and dentary flipped, and frontal shown in dorsal view (modified from Azuma et al., 2016).</td></tr></tbody></table><div class="separator" style="clear: both; text-align: center;"></div><br />An almost complete vertebral series is present.&nbsp; The description states "ten cervical vertebrae are preserved, missing at least the atlas but the materials list only says "eight cervical vertebrae" are present.&nbsp; Hmm.&nbsp; The text states "the middle cervical vertebrae have a highly modified hyposphene-hypantrum with the hypantrum extending ventrally below the dorsal border of the neural canal (Fig. 6b)", but the figure shows nothing extending ventrally into the neural canal and I thought hyposphene-hypantrum articulations were absent from most theropod anterior dorsals, let alone cervicals.&nbsp; "Dorsal centra are longer anteroposteriorly than tall dorsoventrally, unlike the dorsal centra of typical predatory theropods."&nbsp; :|&nbsp; Tell that to coelophysoids, compsognathid-grade taxa, troodontids, microraptorians...&nbsp; "Likely pleurocoels are present in all dorsal vertebrae in the form of longitudinal fossae on the lateral surfaces of centra."&nbsp; Haven't we progressed since the days of Osmolska et al. and Welles calling central fossae pleurocoels?&nbsp; This is another character coded as dromaeosaurid-like, but the posterior dorsals (at least) actually lack pleurocoels as is standard for theropods.&nbsp; Azuma et al. also state "the parapophyses of the dorsal vertebrae including the posterior ones are stalk-like as in derived alvarezsauroids and dromaeosaurids, though they are not as prominent as in the latter groups", but they're actually short.&nbsp; My matrix uses a ratio of parapophyseal length (measured from the ventromedial corner to the apex) and centrum width, so that even <i>Alxasaurus </i>and some <i>Sinraptor </i>and <i>Allosaurus </i>vertebrae count as apomorphic (contra the unchanging TWG matrix), but <i>Fukuivenator </i>still falls short.&nbsp; The authors state "the [sacral] zygapophyses are fused to each other to form a platform lateral to the neural spines, a feature also known in dromaeosaurids", but the figured sacrum lacks neural arches in the first four vertebrae.&nbsp; Azuma et al. say the sacral ribs+diapophyses are "bifurcated distally to contact the ilium, a feature previously unreported in any other theropod", but this is true in e.g. the middle four sacrals of <i>Gallimimus</i>.&nbsp; They also say "the most unusual feature is that the prezygapophyses of the middle caudal vertebrae are distally bifid (Fig. 6i), which has not been reported in any dinosaurs", but this is a standard dromaeosaurid character reported in e.g. <i>Deinonychus </i>(Ostrom, 1969) and <i>Velociraptor </i>(Norell and Makovicky, 1999).<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://3.bp.blogspot.com/-gd99hESPAbU/Vs5g-q2URfI/AAAAAAAAA0w/OZ4YfoirLMY/s1600/Fukuivenator%2Bpes.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="259" src="https://3.bp.blogspot.com/-gd99hESPAbU/Vs5g-q2URfI/AAAAAAAAA0w/OZ4YfoirLMY/s320/Fukuivenator%2Bpes.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Pes of <i>Fukuiraptor paradoxus</i> holotype, as shown in figure 7h on left, and with my reidentification of the phalanges on right (modified from Azuma et al., 2016).&nbsp; Note the longer phalanges on digit II as in other theropods, and how digit II isn't particularly developed into a sickle-claw.</td></tr></tbody></table>Contra the text and coding, the coracoid is proximodistally shallow, unlike pennaraptorans.&nbsp; Surely, figure 7's caption is incorrect and the humeri shown are a right in anterior view and left in lateral view, not the left "in lateral (left) and posterior (right) view."&nbsp; Also, the femur in figure 7f is in medial and posterior views, not lateral and posterior.&nbsp; In figure 7h, the pedal phalanges are almost certainly placed incorrectly, with II-1 and II-2 switched with IV-1 and IV-2, which explains why digit II looks ridiculously short (see figure above).&nbsp; "IV-1" has the standard shape of II-1 with the medial side highly concave (so it actually belongs on the other foot), plus I don't think any terrestrial Mesozoic theropod has IV-1 longer than II-1.&nbsp; The measurement table is partly inconsistent as it has IV-2 subequal to IV-3 and IV-4 in length, unlike the figure.&nbsp; Contra the text and coding, I don't think the second pedal digit looks particularly deinonychosaurian- <i>Tanycolagreus </i>has the same dorsally prominent distal articular surface on II-1, and the ungual in <i>Ornitholestes </i>is comparatively larger.&nbsp; Alas, the text continues to use the II-III-IV manual digit identification, thanks to Xu, which at least Choiniere has disavowed (in his recent <i>Ceratosaurus </i>forelimb paper with Carrano).&nbsp; It was an intriguing idea Xu, but it's long past time to let it die.<br /><br />Azuma et al. add <i>Fukuivenator </i>to Turner et al.'s (2012) TWG analysis, <a href="http://theropoddatabase.blogspot.com/2012/08/turner-et-al-2012-great-review-sloppy.html" target="_blank">which is unfortunately flawed</a> by having numerous taxa left uncoded for entire sections of the matrix, and most of the codings retained from the original TWG matrix of Norell et al. (2001).&nbsp; So don't put much credit into e.g. the joining of oviraptorosaurs and therizinosaurs, because the latter are only represented by the 2001 codings for <i>Alxasaurus</i>, <i>Segnosaurus </i>and <i>Erlikosaurus</i>.&nbsp; No <i>Beipiaosaurus</i>, let alone <i>Falcarius </i>or <i>Jianchangosaurus</i>.&nbsp; In any case, contra the text, they do not recover <i>Fukuivenator </i>as a maniraptoran.&nbsp; In their trees, it's actually in a polytomy with compsognathids, <i>Ornitholestes</i>, ornithomimosaurs and maniraptorans.&nbsp; They also didn't find "<i>Anchiornis </i>and <i>Xiaotingia </i>outside the Troodontidae", instead the base of Deinonychosauria is unresolved with those genera part of the polytomy.&nbsp; Frustratingly, the authors never try excluding taxa a posteriori, so we don't know if e.g. the underlying structure of Turner's tree is still there and <i>Fukuivenator </i>can fall out in multiple places, or if it creates an actual polytomy between some taxa there.&nbsp; Running the matrix myself, I find the latter is true- <i>Fukuivenator</i>'s character combination creates a polytomy there even if it's pruned from the tree a posteriori.&nbsp; Also, the <i>Anchiornis</i>+<i>Xiaotingia </i>situation exists because in a minority of trees these are sister to microraptorians+eudromaeosaurs instead of being basal troodontids.&nbsp; Interestingly, the authors do check how parsimonious it is to place <i>Fukuivenator </i>in alternative positions and find it only takes three more steps to make it a paravian, deinonychosaur or dromaeosaurid.&nbsp; <br /><br />Adding <i>Fukuivenator </i>to the Lori matrix recovers it in a trichotomy with Ornithomimosauria and Maniraptora.&nbsp; Makes sense considering its general basal coelurosaur grade anatomy.&nbsp; There are a few dromaeosaurid-like characters- the fossa around the maxillary fenestra, the short ventral postorbital process, the squamosal shelf over its ventral process, the twisted paroccipital processes, the reduced crista prootica, the bifurcated caudal prezygapophyses.&nbsp; But the vast majority are quite unlike paravians, and as seen above, many of the supposedly dromaeosaurid-like characters aren't present.&nbsp; <a href="http://theropoda.blogspot.com/2016/02/il-paradossale-fukuivenator.html" target="_blank">Andrea Cau recovered <i>Fukuivenator </i>as sister to Pennaraptora in his Megamatrix</a>, which isn't far from my result as only alvarezsauroids and therizinosaurs are between the two <i>Fukuivenator </i>positions, basal members of which (e.g. <i>Haplocheirus</i>, <i>Falcarius</i>) are like <i>Ornitholestes</i>-grade coelurosaurs in much of their anatomy.&nbsp; <br /><br /><b>References</b>- Ostrom, 1969. Osteology of <i>Deinonychus antirrhopus</i>, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History. 30, 165 pp. <br /><br />Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of <i>Velociraptor mongoliensis</i>. American Museum Novitates. 3282, 45 pp.<br /><br />Anonymous, 2009. [3rd new species? Small-sized meat diet dinosaur to restoration Fukui] msn.com 3/18/2009 <a href="http://sankei.jp.msn.com/science/science/090318/scn0903182113002-n1.htm">http://sankei.jp.msn.com/science/science/090318/scn0903182113002-n1.htm</a>&nbsp; <br /><br />Shibata and Azuma, 2010. New dinosaurs from the Lower Cretaceous Kitadani Formation of the Tetori Group, Fukui, Central Japan. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 163A-164A.<br /><br />Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.<br /><br />Azuma, Xu, Shibata, Kawabe, Miyata and Imai, 2016. A bizarre theropod from the Early Cretaceous of Japan highlighting mosaic evolution among coelurosaurians. Scientific Reports. 6, 20478. Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com1tag:blogger.com,1999:blog-3248412803814730250.post-8329584400666060352016-02-18T01:14:00.001-08:002016-02-18T01:14:16.563-08:00Sauropod ThoughtsSo this month I've been concentrating on adding sauropods to the Database, going in reverse chronological order.&nbsp; I find sauropods interesting because unlike theropods, their phylogenetics are almost completely unresolved.&nbsp; I mean sure there's a basic outline and diplodocoids are pretty well researched, but there are huge swaths of cetiosaur-grade and titanosaur(iform/oid) grade taxa that can go almost anywhere between <i>Vulcanodon </i>and Neosauropoda on the one hand, and between <i>Camarasaurus </i>and Eutitanosauria on the other.&nbsp; What follows are some thoughts on taxa or papers describing them...<br /><br /><i>Zby </i>is based on mostly a forelimb that was originally assigned to <i>Turiasaurus </i>in a SVP abstract (Mateus, 2009), but later described as a new genus (Mateus et al., 2014).&nbsp; But the seven characters listed by the latter authors as distinguishing the taxa are only determinable in their holotypes, not the referred specimens of <i>Turiasaurus</i>.&nbsp; Given their stratigraphic and geographic proximity and the variation seen within theropod species, it seems plausible Mateus was right in 2009.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://3.bp.blogspot.com/-mQsqGQWdHe8/VsWGPby6XzI/AAAAAAAAA0A/dF1TRfldBng/s1600/Zby.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="180" src="https://3.bp.blogspot.com/-mQsqGQWdHe8/VsWGPby6XzI/AAAAAAAAA0A/dF1TRfldBng/s320/Zby.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Zby atlanticus</i> (ML 368) (after Mateus et al., 2014).</td></tr></tbody></table><br /><br /><br />I didn't realize how sad the descriptive situation is for <i>Galeamopus</i>. Sure the type skull was described almost a century ago, and the other skulls have featured in discussions of <i>Diplodocus </i>crania, but the postcrania of the type or the new Howe Quarry specimen (SMA 0011) have never been described.&nbsp; We need not only a description of these, but also a paper describing all the supposed <i>Diplodocus </i>skulls, since according to Tschopp et al. (2015), no skulls can actually be referred to that genus.&nbsp; Thus the <i>Diplodocus </i>OTUs in every other analysis are composites.<br /><br />Despite having four generally stellar coauthors, the description of <i>Abydosaurus </i>(Chure et al., 2010) is... well... bad.&nbsp; Even though it has seven supplementary documents, one of which is a spreadsheet of several hundred sauropodomorph tooth elongation indices, there's no measurement table for anything except two cervicals.&nbsp; The most we get for the skull is "The four known skulls of <i>Abydosaurus </i>are nearly the same size, measuring approximately 0.5 m long and half as tall posteriorly."&nbsp; You could have used less words and given us precise data for all four specimens.&nbsp; And the table with the cervical measurements includes data for seven other sauropods, all but two of which are from previously published literature anyway.&nbsp; Why spend space on data already out there instead of publishing new data?&nbsp; The referred postcrania don't have repository numbers listed either.&nbsp; Finally, the phylogenetic analysis goes out of its way to delete data from Wilson's original version, removing all but two non-neosauropods, combining genera to make four suprageneric OTUs, deleting characters that don't vary due to those removals, etc..&nbsp; All this does is make it more tedious for future workers to add <i>Abydosaurus </i>to the base version of Wilson's matrix that tens of other papers use, and it's not like Wilson's 234 character, 27 taxon matrix is unwieldy for any 2010 computer to handle.&nbsp; They just spent a lot of time and effort to make the analysis worse.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="https://2.bp.blogspot.com/-0gben0twDYo/VsWHQmmvjcI/AAAAAAAAA0I/vn8BmsTA78c/s1600/Normanniasaurus.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="106" src="https://2.bp.blogspot.com/-0gben0twDYo/VsWHQmmvjcI/AAAAAAAAA0I/vn8BmsTA78c/s320/Normanniasaurus.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Proximal caudal vertebra of <i>Normanniasaurus genceyi</i> (MHNH-2013.2.1) (after Le Loeuff et al., 2013). Scale = 10 cm.</td></tr></tbody></table><br /><i>Normanniasaurus </i>(Le Loeuff et al., 2013) is one of the most fragmentary taxa I've seen erected in the past few decades (the holotype consists of two presacral prezygapophyses, three fused sacral centra, an incomplete proximal caudal vertebra, an incomplete mid caudal vertebra, partial scapula, two ilial fragments, ischia [one partial, one fragmentary], a femoral fragment and a fibular fragment).&nbsp; This is quite ironic considering Le Loeuff's famous 1993 paper declaring most European titanosaurs to be indeterminate (<i>Macrurosaurus</i>, <i>Hypselosaurus</i>, <i>Aepisaurus</i>, <i>"Titanosaurus" lydekkeri</i>).&nbsp; Most of the supposedly diagnostic characters listed by the authors are either primitive for titanosauriforms (presacral vertebrae with hyposphene-hypantrum articulation; middle caudal vertebrae amphicoelous; ilium with anterolaterally expanded blade) or classic characters diagnosing larger clades within it (internal texture of presacral vertebrae camellate; proximal caudal vertebrae deeply procoelous; middle caudal vertebrae with anteriorly placed neural arch).&nbsp; I'm very doubtful whether the remaining few caudal characters will prove diagnostic if anyone looks into them.&nbsp; When added to Mannion et al.'s (2013) matrix, <i>Normanniasaurus </i>resolves as a eutitanosaur closer to <i>Saltasaurus </i>than opisthocoelicaudiines.&nbsp; <br /><br />Finally, Mannion and Otero (2012) did a superb job redescribing <i>Argyrosaurus</i>.&nbsp; The taxon is only known from a forelimb, though apparently the entire skeleton was originally there but did not survive excavation.&nbsp; This is my favorite kind of paper- redescribing a taxon known for decades in a modern context.&nbsp; Mannion's one of my favorite sauropod authors, having redescribed <i>Mongolosaurus </i>(2010), <i>"Brachiosaurus" atalaiensis</i> (2013) and <i>B? nougaredi</i> (2013).&nbsp; Mannion and Otero didn't include <i>Argyrosaurus </i>in a phylogenetic analysis, saying it has some plesiomorphic characters despite sharing many characters with derived titanosaurs.&nbsp; I coded <i>Argyrosaurus </i>in Mannion et al.'s macronarian matrix, and I have to say the characters are well formed, so that coding is objective.&nbsp; I'm not a sauropod expert, so I don't have the background comprehension of sauropod characters, but Mannion makes them easy to code.&nbsp; <i>Argyrosaurus </i>ends up as a nemegtosaurid (sister to <i>Mongolosaurus </i>and <i>Rapetosaurus</i>), so despite Mannion et al.'s lack of many lithostrotians, the genus seems to be a member of the clade.&nbsp; If it is a nemegtosaurid, Argyrosauridae Bonaparte 1987 has priority over Nemegtosauridae Upchurch 1995.&nbsp; Interesting.&nbsp; One critique of the paper is that it doesn't specify the horizon and locality of referred <i>Argyrosaurus </i>specimens (so I had to refer back to Powell 2003 and Bonaparte and Gasparini 1979), and proposes these are nomina dubia without detailed examination or comparison.<br /><br />Will the next post get back to theropods? Depends on the papers published in the near future.&nbsp; Come on, O'Connor, I dare ya. ;)<br /><br /><b>References</b>- Mateus, 2009. The sauropod dinosaur <i>Turiasaurus riodevensis</i> in the Late Jurassic of Portugal. Journal of Vertebrate Paleontology. 29(3), 144A.<br /><br />Chure, Britt, Whitlock and Wilson, 2010. First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition. Naturwissenschaften. 97(4), 379-391.<br /><br />Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod dinosaur <i>Argyrosaurus superbus</i>, with a description of a new titanosaur genus. Journal of Vertebrate Paleontology. 32(3), 614-638.<br /><br />Le Loeuff, Suteethorn and Buffetaut, 2013. A new sauropod dinosaur from the Albian of Le Havre (Normandy, France). Oryctos. 10, 23-30.<br /><br />Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese sauropod dinosaur <i>Lusotitan atalaiensis</i> (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of the Linnean Society. 168(1), 98-206. <br /><br />Mateus, Mannion and Upchurch, 2014. <i>Zby atlanticus</i>, a new turiasaurian sauropod (Dinosauria, Eusauropoda) from the Late Jurassic of Portugal. Journal of Vertebrate Paleontology. 34(3), 618-634.<br /><br />Tschopp, Mateus and Benson, 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ. 3:e857.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com3tag:blogger.com,1999:blog-3248412803814730250.post-43341674373711129032016-01-23T23:59:00.000-08:002016-01-23T23:59:02.276-08:00Dracoraptor thoughtsSo the Welsh coelophysoid has been published, with the dudebro name of <i>Dracoraptor</i>.&nbsp; There seems to have been a lack of rigorous editing, as exemplified by "<i>Podekosaurus</i>", <i>Eshanosaurus </i>being a therizinosaurid (as opposed to -oid or -ian), BMNH and NHMUK being used in the same sentence for specimens, and that's just the first two pages.&nbsp; I also think the kind of title this paper has is silly ("The oldest Jurassic dinosaur") since it just specifies position within a known range.&nbsp; It's like having "The smallest dinosaur under two meters long" or "the westernmost dinosaur found in China." Pluses are it's in open access and the photos are color with great resolution.<br /><br />As for the actual description, there's a discrepancy in the elements that are shown in the skeletal reconstruction (fig. 5), listed under the holotype, listed at the beginning of each anatomical area, and actually described.&nbsp; For instance, figure 5 shows a squamosal, angular and articular as preserved (and not just as molds or tentatively identified), which are listed under holotype.&nbsp; Yet only "more caudal elements of the lower jaws" are listed under the cranial elements heading, and none of these elements are described.&nbsp; But a ?nasal is described and figured, but not in figure 5 or noted in the cranial intro.&nbsp; In total it seems more written by committee than cohesive.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-hb0XHzvzCYs/VqHn0d-47AI/AAAAAAAAAzo/0Y6OKKUQInM/s1600/Dracoraptor%2Binterp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://3.bp.blogspot.com/-hb0XHzvzCYs/VqHn0d-47AI/AAAAAAAAAzo/0Y6OKKUQInM/s320/Dracoraptor%2Binterp.jpg" width="296" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Color figures- holotype of <i>Dracoraptor hanigani</i> (NMW 2015.5G.3) with: A- supposed manual ungual III, B- supposed manual ungual II, C- supposed manual ungual I, D- supposed furcula (after Martill et al., 2016).&nbsp; Black and white figures from left to right- pedal unguals I and II of <i>Coelophysis bauri</i> NMMNH P-42200 (after Rinehart et al., 2009); pedal ungual II in dorsal view of <i>Liliensternus liliensterni</i> syntype MB.R.2175 (after Huene, 1934); metacarpal II of <i>Coelophysis? rhodesiensis</i> holotype QG 1 (after Galton, 1971); furcula of <i>kayentakatae </i>holotype MNA V2623 in dorsal and anterior views (after Tykoski et al., 2002); ninth dorsal rib of <i>Allosaurus fragilis</i> USNM 8367 (after Gilmore, 1920). Red lines indicate element identifications I propose for <i>Dracoraptor</i>. </td></tr></tbody></table>Note the maxillae are in medial view, as the figure caption states, but contra the text.&nbsp; Contra the text and reconstruction, only the small anteriormost jugal groove probably articulated with the maxilla.&nbsp; The longer ridge and groove on the main body is common in early dinosaurs (e.g. <i>Herrerasaurus</i>) and non-articulating.&nbsp; The supraoccipital is in anterior (internal) view, not ventral view as stated by the text.&nbsp; Note the large posttemporal fenestrae as in <i>Silesaurus </i>but unlike dinosaurs.&nbsp;<br /><br />The cervical is not opisthocoelous, contra the text, as the anterior central surface is slightly concave.&nbsp; The supposed first caudal is near certainly a ?last sacral based on the broad transverse processes originating on the centrum (compare to e.g. <i>Staurikosaurus</i>).&nbsp; The authors do call it "a partially sacralised element", but any ambiguity seems unnecessary.&nbsp; The next element could easily be a sacral too, though its more fragmented condition makes this more uncertain.&nbsp; I'm doubtful the supposed furcula is correctly identified.&nbsp; One side is much narrower than the other, and each is curved in a different direction (thin side concave toward the outside of the angle).&nbsp; Furcular arms are subequal in width, and those of coelophysoids (e.g. <i>kayentakatae</i>) are basically circular in section, so that twisting in <i>Dracoraptor </i>is not an excuse.&nbsp; It's more probably a posterior dorsal rib, which are also similar in having a ridge along the outside corner. The tuberculum may be covered in matrix.&nbsp;<br /><br />Amusingly, Martill et al. state that on the humerus "a suture between the articulatory epiphyseal surfaces is well defined."&nbsp; And for the ulna, "the proximal condyle surface is smooth and well defined, marked from the diaphysis by a prominent suture."&nbsp; Dinosaurs aren't mammals, people.&nbsp; They don't have separately ossifying epiphyses.&nbsp; For the femur, "the femoral head (greater trochanter) is "hooked'."&nbsp; Er, those are very different structures, not synonyms.&nbsp; The authors say "A calcaneum is not present. Two distal tarsals (dt III &amp; dt IV) and part of a putative third are present in a row."&nbsp; I'm pretty sure no archosauriforms actually have three distal tarsals per pes (certainly no theropods do), so that 'putative third' is more likely the supposedly missing calcaneum, especially as it's placed right next to distal tarsal IV (confusingly labeled 'Ldtii').&nbsp; <br /><br />The supposed "?Metaacarpal of digit I" [sic] is a metacarpal II, very similar to <i>rhodesiensis</i>, more elongate than metacarpal I and more robust than metacarpal III.&nbsp; Even <a href="https://pterosaurheresies.wordpress.com/2016/01/22/dracoraptor-a-scrappy-new-earliest-jurassic-theropod-from-wales/" target="_blank">Peters got this right in his reinterpretation</a>, which is especially sad for Martill et al..&nbsp; While I haven't identified all of the phalanges in this block, it's clear Martill et al.'s statement "they are assumed to be from the left manus as they are associated with the left radius and ulna" is in error.&nbsp; For instance, the phalanx underlying the proximal radius is too large to belong to any manual digit and is probably pedal phalanx III-1, while supposed manual unguals I and III lack flexor tubercles ("I" shows an obvious depression in that area) and at least "III" is virtually straight.&nbsp; These unguals more nearly match pedal unguals of e.g. <i>Coelophysis </i>and <i>Liliensternus</i>, while supposed ungual II is manual due to its curvature and large flexor tubercle.&nbsp; Among other phalanges, that at the distal end of metacarpal II matches a manual phalanx II-1, that on the proximal end of metacarpal II belongs to manual digit III, that between unguals "I" and "II" looks like its from pedal digit II, and the small one by the anterior end of the dorsal centrum would be manual IV-1.&nbsp; The latter suggests a less reduced digit IV than coelophysids or <i>Herrerasaurus</i>.&nbsp; Thus the manual reconstruction with its short penultimate phalanges and metacarpal ratios should not be trusted.<br /><br />The phylogenetic analysis is&nbsp; based on Nesbitt et al.'s <i>Tawa </i>matrix.&nbsp; One good thing is that Martill et al. split Nesbitt's composite characters, which had the effect of Nesbitt assuming non-homology for e.g. subnarial foramina in different positions, jugal ridges of varying sharpness, etc..&nbsp; A negative point is that Martill et al. do not order any characters "because it assumes a complete fossil record and a direction to evolution."&nbsp; But ordering does not assume these, it merely observes that some states are more similar to others.&nbsp; So e.g. if a taxon has 5 premaxillary teeth, it's more similar to a taxon with 6 teeth than to one with 3 teeth.&nbsp; Ironically, some of Martill et al.'s states imply ordering, such as their "6+ premaxillary teeth" state, which only makes sense if taxa with 6 teeth are more similar to taxa with 7, 8, etc. teeth than to those with less than six teeth.&nbsp; They recover <i>Dracoraptor </i>as the basalmost coelophysoid, sister to Coelophysidae. I like how they note "an unnamed clade containing <i>Velociraptor</i>, <i>Allosaurus </i>and <i>Piatnitzkysaurus</i>."&nbsp; Ah, you mean that obscure group called Tetanurae. ;)<br /><br />Luckily for me, I have a version of Nesbitt et al.'s matrix on hand with characters properly ordered, and basically every basal dinosauriform added (as used in <a href="http://theropoddatabase.blogspot.com/2015/06/chilesaurus-brings-out-bandit-in-me.html" target="_blank">my <i>Chilesaurus</i> post</a>).&nbsp; I've also made a ton of corrections, though have by no means gone through the whole thing yet.&nbsp; I did go through Martill et al.'s <i>Dracoraptor </i>codings though, and 15% are miscoded.&nbsp; These range from consequences of my reidentifications above, to general errors (e.g. while coded as lacking a promaxillary fenestra, you can't tell that as the maxillae are in medial view), to times where I don't think the authors understood the character (e.g. it's coded as having an anterior hollow on the astragalus, but as used by Nesbitt this is a feature that has been lost in all avemetatarsalians).&nbsp; After correcting <i>Dracoraptor</i>'s codings, it emerges as sister to Neotheropoda (Ceratosauria+Tetanurae; Averostra of some authors).&nbsp; But Neotheropoda has an odd topology where <i>Chilesaurus </i>is sister to <i>Velociraptor</i>, then <i>Ceratosaurus</i>, <i>Piatnitzkysaurus </i>and <i>Allosaurus </i>form a clade.&nbsp; Because this makes the basal condition of Neotheropoda very different than the consensus, I deleted the highly modified <i>Velociraptor</i>.&nbsp; Recall from the <i>Chilesaurus</i> post that <i>Velociraptor </i>only clades with <i>Piatnitzkysaurus </i>and <i>Allosaurus </i>in Nesbitt's uncorrected trees because it's miscoded for 24% of the characters that make it seem more like a "normal" theropod.&nbsp; After Velociraptor's deletion Neotheropoda has the standard topology, and <i>Dracoraptor </i>is sister to a <i>Daemonosaurus</i>+<i>Chilesaurus </i>clade, which is itself sister to Avepoda (Neotheropoda of some authors).&nbsp; This is rather like <a href="http://theropoda.blogspot.com/2016/01/dracoraptor-il-piu-antico-neotheropode.html" target="_blank">Cau's result</a>, where <i>Dracoraptor </i>grouped with <i>Daemonosaurus </i>and <i>Tawa</i>.&nbsp; In my trees, <i>Tawa </i>is the next taxon stemward.&nbsp; Because <i>Chilesaurus</i>' basal theropod status is my pet theory and the taxon is also highly modified, I tried deleting it as well.&nbsp; The result then is that <i>Dracoraptor </i>forms a polytomy with <i>Tawa</i>, <i>Daemonosaurus</i>, Coelophysidae and more derived theropods.<br /><br />Thus it seems Cau's and my matrices agree in placing <i>Dracoraptor </i>by <i>Daemonosaurus</i>.&nbsp; Note both share three premaxillary teeth (also in <i>Chilesaurus </i>and <i>Tawa</i>), short snouts with few maxillary teeth (also in <i>Chilesaurus</i>, not in <i>Tawa</i>), a lacrimal flange that does not overlap the antorbital fenestra (not in <i>Tawa</i>; unknown in <i>Chilesaurus</i>), and elongate cervicals with single pleurocoels (also in <i>Tawa</i>; elongate centra but double pleurocoels in <i>Chilesaurus</i>).&nbsp; <br /><br /><b>References</b>- Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera <i>Antrodemus</i> (<i>Allosaurus</i>) and <i>Ceratosaurus</i>. Bulletin of the United States National Museum. 110, 1-154. <br /><br />Huene, 1934. Ein neuer Coelurosaurier in der thüringischen Trias [A new coelurosaur in the Thuringian Trias]. Paläontologische Zeitschrift. 16(3/4), 145-170.<br /><br />Galton, 1971. Manus movements of the coelurosaurian dinosaur <i>Syntarsus </i>and opposability of the theropod hallux. Arnoldia. 5(15), 1-8.<br /><br />Tykoski, Forster, Rowe, Sampson and Munyikwa, 2002. A furcula in the coelophysid theropod <i>Syntarsus</i>. Journal of Vertebrate Paleontology. 22(3), 728-733. <br /><br />Rinehart, Lucas, Heckert, Spielmann and Celeskey, 2009. The Paleobiology of <i>Coelophysis bauri</i> (Cope) from the Upper Triassic (Apachean) Whitaker quarry, New Mexico, with detailed analysis of a single quarry block. New Mexico Museum of Natural History and Science Bulletin. 45, 260 pp.<br /><br />Martill, Vidovic, Howells and Nudds, 2016. The oldest Jurassic dinosaur: A basal neotheropod from the Hettangian of Great Britain. PLoS ONE. 11(1), e0145713. Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com8tag:blogger.com,1999:blog-3248412803814730250.post-40804236476685320892016-01-15T02:09:00.001-08:002016-01-16T03:42:30.211-08:00Nanotyrannus was a groovy babyThe title works best when read in Austin Powers' voice. <br /><br />So a new paper's out, and let's just say it's not very good.&nbsp; Schmerge and Rothschild's (2016) premise is that <i>Nanotyrannus</i> has a lateral dentary groove, so based on this and adding it to Brusatte et al.'s (2010) tyrannosauroid matrix, it's an albertosaurine instead of a tyrannosaurine, let alone a juvenile <i>Tyrannosaurus</i>.<br /><br />The first problem is that the lateral dentary groove has never been a great character, since its depth varies so much between taxa that numerous genera have been coded both ways in different TWG analyses.&nbsp; I would say for instance that the slight longitudinal depression in <i>Dryptosaurus </i>(Brusatte et al., 2011; fig. 3A) doesn't qualify, contra the authors.&nbsp; Indeed, Carr (online) list additional tyrannosauroids he interprets as having the groove, which were not coded that way by Brusatte et al..&nbsp; Of these, I'd agree with <i>Raptorex</i>, and the juvenile <i>Bistahieversor </i>and <i>Tarbosaurus</i>, but view e.g. the <i>Bistahieversor </i>type and <i>Alioramus </i>(<i>altai</i>'s type) to be more like <i>Dryptosaurus</i>.&nbsp; Thus I agree with Headden (DML, 2016) that the character shows ontogenetic variation.&nbsp; But my main point is that it's so subjective that authors can't agree on which morphologies count, and that any attempt to quantify depth and continuity (many individuals have grooves that vary in depth along the dentary, most obviously visible in the CT scan of <i>Tarbosaurus </i>juvenile IGM 107/7) won't be easy.<br /><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-DmJyTIvzq1s/Vpi1yChbp_I/AAAAAAAAAyI/bsfSgzrPp-I/s1600/tyrannosauroid%2Bdentary%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="250" src="http://1.bp.blogspot.com/-DmJyTIvzq1s/Vpi1yChbp_I/AAAAAAAAAyI/bsfSgzrPp-I/s320/tyrannosauroid%2Bdentary%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Dentaries of tyrannosauroid taxa (some flipped) showing degree of lateral groove.&nbsp; Left (top to bottom)- <i>Bistahieversor </i>juvenile paratype NMMNH P-25049 (after Carr and Williamson, 2010); <i>Tarbosaurus </i>juvenile IGM 107/7 (after Tsuihiji et al., 2011); <i>Nanotyrannus </i>BMRP 2002.4.1 'Jane' (after Schmerge and Rothschild, 2016); <i>Raptorex </i>juvenile holotype LH PV18 (after Sereno et al., 2009). Right (top to bottom)- <i>Dryptosaurus </i>holotype ANSP 9995 (after Brusatte et al., 2011); <i>Bistahieversor </i>holotype NMMNH P-27469 (after Carr and Williamson, 2010); <i>Alioramus </i>IGM 100/1844 (after Brusatte et al., 2012); <i>Tyrannosaurus </i>FMNH PR2081 (after Brochu, 2003).&nbsp; I'd say those on the left have a strong groove while those on the right lack one, and the least <i>Dryptosaurus </i>has less of a groove than e.g. <i>Bistahieversor </i>or <i>Tyrannosaurus</i>, contra Schmerge and Rothschild's recoding of it.&nbsp; Also note deeper grooves are common in juvenile tyrannosaurids.</td></tr></tbody></table><br />The second problem is that even if Schmerge and Rothschild had an unassailable, quantifiable character difference between <i>Nanotyrannus </i>and <i>Tyrannosaurus </i>that was not known to be ontogenetically variable, it's STILL just one character.&nbsp; They're being BANDit-esque in assuming one character can override the totality of character evidence.&nbsp; Sure they go on to list five other characters supposedly shared between <i>Nanotyrannus </i>and albertosaurines, but not only are some obviously plesiomorphies&nbsp; (maxillary fenestra placed posteriorly; rounder orbit; greater dentary tooth count), the authors actually got the maxillary fenestra character backwards (they claim "contact of the maxillary fenestra with the rostral margin of the antorbital fossa" unites <i>Nanotyrannus </i>and albertosaurines).&nbsp; The juvenile <i>Tarbosaurus </i>and <i>Maleevosaurus</i>' type are sufficient to show it's expected in juvenile <i>Tyrannosaurus </i>anyway.&nbsp; What about the cladistic analysis, you ask?&nbsp; While I haven't seen the matrix yet<b>*</b>, I'm betting the authors coded <i>Nanotyrannus </i>as if it was an adult.&nbsp; And we already know juvenile tyrannosaurines emerge more basally when coded as adults.&nbsp; Carr (2005) found <i>Nanotyrannus</i>' holotype to be sister to <i>Daspletosaurus</i>+<i>Tarbosaurus</i>+<i>Tyrannosaurus</i>, and the <i>Stygivenator </i>holotype to be "in a polytomy consisting of <i>Teratophoneus </i>and all non-tyrannosaurine tyrannosauroids."&nbsp; Tsuihiji et al. (2011) found their <i>Tarbosaurus </i>juvenile to emerge sister to albertosaurines+tyrannosaurines.&nbsp; So this is an unsurprising result even assuming Schmerge and Rothschild's coding was competent.<br /><br /><b>Edit- the matrix is terribly undercoded. &nbsp; Only one character of 27 coded for the nasal and lacrimal, the squamosal and quadratojugal are completely uncoded, the entire skull roof, palate and braincase uncoded, postdentary mandibular bones uncoded, only one postcranial character coded.&nbsp; Did nobody check the matrix and notice the huge swath of question&nbsp; marks?!&nbsp; Seriously, Cretaceous Research, this is just sad.</b><br /><br />Finally, the authors don't understand how to compute character changes, when they compare the steps necessary to move <i>Nanotyrannus </i>to <i>Tyrannosaurus</i>' sister OTU.&nbsp; Note we're never presented with the actual number of steps such a move adds to the analysis, let alone told if that number is statistically significant.&nbsp; Instead, Schmerge and Rothschild just show the distribution of their dentary groove character if the move is made.&nbsp; Even accepting their coding is correct, the correct answer is that moving <i>Nanotyrannus </i>to <i>Tyrannosaurus </i>would add a single step, namely <i>Nanotyrannus </i>evolving that character in convergence with albertosaurines.&nbsp; But the authors claim "this tree requires 5 more independent losses of the dentary groove than the tree proposed in this study."&nbsp; Wha?!&nbsp; How do they justify this?&nbsp; Because for some never explained reason, the authors assume the groove is present along the entire 'backbone' of the cladogram, so that every tyrannosauroid that lacks the groove independently loses it.&nbsp; How terribly unparsimonious!&nbsp; This takes 11 steps in their tree 3B, whereas the most parsimonious phylogenetic reconstruction would take only four steps assuming the groove is present in the outgroup as the authors believe.&nbsp; All that's necessary is the loss of the groove for taxa more derived than proceratosaurids, then independent development in <i>Dryptosaurus</i>, albertosaurines and <i>Nanotyrannus</i>.&nbsp; I can only suppose the authors view the groove as being incapable of parallel development.&nbsp; It's just formed by a branch of the (?)trigeminal nerve being more depressed into the dentary's lateral surface, hardly the "dramatic change (e.g., metamorphosis) [that] would need to be invoked to explain the loss of this feature through maturation" that Schmerge and Rothschild claim.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-lKd_vwDDjAY/VpjDkQh9bHI/AAAAAAAAAyc/g7l5inzcw9Y/s1600/tyranno%2Bgroove%2Bphylo.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://4.bp.blogspot.com/-lKd_vwDDjAY/VpjDkQh9bHI/AAAAAAAAAyc/g7l5inzcw9Y/s320/tyranno%2Bgroove%2Bphylo.jpg" width="214" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Cladogram of tyrannosauroids with deep lateral dentary grooves indicated with bold lines.&nbsp; Middle- figure 3B from Schmerge and Rothschild (2016) showing their proposed evolution of dentary grooves.&nbsp; Top- number of steps assumed by Schmerge and Rothschild, with steps indicated by circles.&nbsp; Bottom- most parsimonious distribution of steps given an outgroup with dentary grooves.</td></tr></tbody></table><br />In conclusion, I don't see how this paper made it to publication.&nbsp; The anatomical structure in question has a controversial distribution, Tsuihiji et al. (2011) destroys the entire premise but is never addressed or referenced, the authors never even consider that coding <i>Nanotyrannus </i>as an adult is problematic (Carr 1999 showed that the holotype has immature bone grain, so even if it's an albertosaurine, it's a juvenile), and they don't understand how character state reconstruction works.&nbsp; Using 'key characters', presumptions about reversability, an ignorance of the recent literature, misunderstanding cladistics... did Schmerge inherit BANDit-style biases working at the University of Kansas?<br /><br /><b>References</b>- Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Theropoda). Journal of Vertebrate Paleontology. 19(3), 497-520. <br /><br />Brochu, 2003. Osteology of <i>Tyrannosaurus rex</i>: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Society of Vertebrate Paleontology Memior. 7, 138 pp. <br /><br />Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. PhD thesis. University of Toronto. 1170 pp.<br /><br />Sereno, Tan, Brusatte, Kriegstein, Zhao and Cloward, 2009. Tyrannosaurid skeletal design first evolved at small body size. Science. 326(5951), 418-422.<br /><br />Carr and Williamson, 2010. <i>Bistahieversor sealeyi</i>, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea. Journal of Vertebrate Paleontology. 30(1), 1-16.&nbsp; <br /><br />Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms. Science. 329, 1481-1485.<br /><br />Brusatte, Benson and Norell, 2011. The anatomy of <i>Dryptosaurus aquilunguis</i> (Dinosauria: Theropoda) and a review of its tyrannosauroid affinities. American Museum Novitates. 3717, 53 pp. <br /><br />Tsuihiji, Watabe, Tsogtbaatar, Tsubamoto, Barsbold, Suzuki, Lee, Ridgely, Kawahara and Witmer, 2011. Cranial osteology of a juvenile specimen of <i>Tarbosaurus bataar</i> (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia. Journal of Vertebrate Paleontology. 31(3), 497-517.<br /><br />Brusatte, Carr and Norell, 2012. The osteology of <i>Alioramus</i>, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History. 366, 197 pp. <br /><br />Carr, online 2016. <a href="http://tyrannosauroideacentral.blogspot.com/2016/01/by-way-that-groove-is-also-seen-in.html">http://tyrannosauroideacentral.blogspot.com/2016/01/by-way-that-groove-is-also-seen-in.html</a><br /><br />Headden, DML 2016. <a href="http://dml.cmnh.org/2016Jan/msg00074.html">http://dml.cmnh.org/2016Jan/msg00074.html</a><br /><br />Schmerge and Rothschild, 2016. Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus <i>Nanotyrannus </i>Bakker et al., 1988. Cretaceous Research. 61, 26-33.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com3tag:blogger.com,1999:blog-3248412803814730250.post-82978757154398439002016-01-12T03:04:00.000-08:002016-01-12T18:12:40.449-08:00How not to redefine OrnithuromorphaThe answer is "any possible way."&nbsp; As in, DON'T redefine Ornithuromorpha Chiappe et al. 1999, which has until now only been given two definitions, both node-based.&nbsp; There's Chiappe's (2001) original "<i>Patagopteryx </i>+ <i>Vorona </i>+ Ornithurae" and his later (2002) more refined "<i>Patagopteryx </i>+ Ornithurae", both based on the same topology, with a trichotomy between <i>Patagopteryx</i>, <i>Vorona </i>and Ornithurae.&nbsp; Ornithurae in this case is Chiappe's version which was <i>Hesperornis </i>plus modern birds.<br /><br />Yet in the description of their new bird <strike><i>Dingornis </i></strike><b>[edit] </b><i><b>Dingavis</b> longimaxilla</i> (first mentioned in Wang et al., 2015), O'Connor et al. (2016) redefine Ornithuromorpha to be stem-based.&nbsp; Their Ornithuromorpha is "The first ancestor of Neornithes that is not also an ancestor of the Enantiornithes, and all of its descendants."<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-9HAUbQvIVEA/VpTR4oyOCkI/AAAAAAAAAx0/XRLGXrxVtSw/s1600/Dingavis.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://3.bp.blogspot.com/-9HAUbQvIVEA/VpTR4oyOCkI/AAAAAAAAAx0/XRLGXrxVtSw/s320/Dingavis.jpg" width="273" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Dingavis longimaxilla</i> IVPP V20284 (scale = 20 mm; after O'Connor et al., 2016).</td></tr></tbody></table><br />First of all, will everyone PLEASE start paying attention to Phylocode Article 11.1- "All specifiers used in node-based and branch-based definitions of clade names, and one of the specifiers used in apomorphy-based definitions of clade names, are species or specimens."&nbsp; Chiappe had the excuse that he was working over a decade ago.&nbsp; But in 2016 that doesn't cut it.&nbsp; And it's SO easy!&nbsp; Just say "<i>Passer domesticus</i> &lt; - <i>Enantiornis leali</i>".&nbsp; Done!<br /><br />Second, O'Connor et al.'s excuse is that "Although the proposed definition does not strictly equate with the published node-based definition, it does provide a formal definition for the current widespread usage of this term in most recent literature (Bell et al. 2010; O’Connor et al. 2010; S. Zhou et al. 2013a; M. Wang et al. 2015)."&nbsp; Let's review those...<br />- Bell et al. (2010) in their description of <i>Hollanda </i>have <i>Patagopteryx </i>as the most basal taxon closer to modern birds than enantiornithines, so have no need for a different definition than Chiappe's and don't explicitly use one either.<br />- O'Connor et al. (2010) in their description of <i>Longicrusavis </i>have the same situation.<br />- Zhou et al. (2013) in their description of new <i>Archaeorhynchus </i>specimens don't even show a topology or mention <i>Patagopteryx</i>.&nbsp; Notably, Zhou and Zhang (2006) in the original description of <i>Archaeorhynchus </i>just deleted <i>Patagopteryx</i> from the matrix they used without justification.&nbsp; Why do you hate <i>Patagopteryx </i>so much, Zhou?<br />- Wang et al. (2015) in their description of new <i>Gansus </i>material do indeed place Ornithuromorpha in the wrong position, as they have <i>Archaeorhynchus </i>and <i>Jianchangornis </i>further from modern birds than <i>Patagopteryx </i>or <i>Vorona</i>, yet include them among Ornithuromorpha.&nbsp; Note O'Connor was second author here.<br />So only one paper listed actually uses Ornithuromorpha as if it were a stem, and that shared a coauthor.&nbsp; Hardly a convincing case that the usage is "widespread."&nbsp; Sure there are other papers that use it this way, but almost all have O'Connor as a coauthor.&nbsp; You don't get to use a name incorrectly tens of times and then say "looks like all the papers are using it this way, we better change it."<br /><br />Third, by redefining names you're countering the entire point of phylogenetic nomenclature.&nbsp; The beauty of the system is that regardless of your topology, you can apply clade names because their definitions stay constant.&nbsp; Just find <i>Passer </i>and <i>Ornithomimus </i>in your tree and you'll always know where Maniraptora goes.&nbsp; But if you pull a Sereno and declare Maniraptora to be "<i>Oviraptor </i>+ <i>Passer</i>" instead, then your concept is no longer the same.&nbsp; Ditto for Ornithuromorpha.<br /><br />"<i>Passer domesticus</i> &lt; - <i>Enantiornis leali</i>" is certainly a clade that deserves a name, but it's not the more exclusive Ornithuromorpha.&nbsp; We didn't call the clade Ornithurae just because the BANDits got it wrong so often in the 90s, and we should aim to be better this time as well.<br /><br /><br /><br />As an aside,&nbsp; O'Connor et al. state "Although we do not consider [<i>Gansus zheni</i>] to be referable to <i>Gansus</i>, previously only known from the Xiagou Formation (You et al. 2006), we also recognize minor differences that suggest they are not referable to <i>Iteravis huchzermeyeri</i>."&nbsp; After <a href="http://theropoddatabase.blogspot.com/2014/12/gansus-zheni-is-iteravis.html" target="_blank">my in depth study</a>, I'm skeptical.&nbsp; Notably, O'Connor et al. did not add <i>zheni </i>to their matrix, although <i>Iteravis </i>emerged in a polytomy with <i>Gansus</i>.&nbsp; <br />Is <i>Dingavis </i>from the same locality another synonym of <i>Iteravis</i>?&nbsp; While it does seem to share the ventrally concave ischium with small mid dorsal process (also in <i>Piscivoravis</i>, <i>Yanornis</i> and <i>Gansus</i>), pedal digit IV is shorter (89% of III excluding unguals; compared to 99-110% in <i>Iteravis</i>), the ectethmoid (lacrimal of O'Connor et al.) is more acute, the carpometacarpus more elongated, and phalanx III-1 lacks the lateral process.&nbsp; So offhand, I'd agree they're distinct.<br /><br /><b>References</b>- Chiappe, 2001. Phylogenetic relationships among basal birds. In Gauthier and Gall (eds). New perspectives on the origin and early evolution of birds: Proceedings of the international symposium in honor of John H. Ostrom. Peabody Museum of Natural History. 125-139. <br /><br />Chiappe, 2002. Basal bird phylogeny: Problems and solutions. In Chiappe and Witmer (eds). Mesozoic birds: Above the heads of dinosaurs. University of California Press. 448-472. <br /><br />Zhou and Zhang, 2006. A beaked basal ornithurine bird (Aves, Ornithurae) from the Lower Cretaceous of China. Zoologica Scripta. 35, 363-373.<br /><br />Bell, Chiappe, Erickson, Suzuki, Watabe, Barsbold and Tsogtbaatar, 2010. Description and ecologic analysis of <i>Hollanda luceria</i>, a Late Cretaceous bird from the Gobi Desert (Mongolia). Cretaceous Research. 31(1), 16-26.<br /><br />O’Connor, Gao and Chiappe, 2010. A new ornithuromorph (Aves: Ornithothoraces) bird from the Jehol Group indicative of higher-level diversity. Journal of Vertebrate Paleontology. 30(2), 311-321.<br /><br />Zhou, Zhou and O'Connor, 2013. Anatomy of the basal ornithuromorph bird <i>Archaeorhynchus spathula</i> from the Early Cretaceous of Liaoning, China. Journal of Vertebrate Paleontology. 33(1), 141-152. <br /><br />Wang, Clarke and Huang, 2015. Ornithurine bird from the Early Cretaceous of China provide new evidence for the timing and pattern of the evolution of avian skull. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 233.<br /><br /><br />Wang, O'Connor, Li and You, 2015. New information on postcranial skeleton of the Early Cretaceous <i>Gansus yumenensis</i> (Aves: Ornithuromorpha), Historical Biology. DOI: 10.1080/08912963.2015.1006217 <br /><br />O'Connor, Wang and Hu, 2016. A new ornithuromorph (Aves) with an elongate rostrum from the Jehol Biota, and the early evolution of rostralization in birds. Journal of Systematic Palaeontology. DOI: 10.1080/14772019.2015.1129518Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-57665773517087450172016-01-06T02:03:00.000-08:002016-01-16T22:58:23.191-08:00Chiappeavis is just another PengornisThe last Mesozoic dinosaur named in 2015 is <i>Chiappeavis magnapremaxillo</i> (O'Connor et al., 2015b), electronically published on December 31st.&nbsp; First of all, what a stupid species name.&nbsp; Second, examination of the supposed diagnostic characters shows that like <i>Parapengornis </i>and IVPP V18632, it's just another example of <i>Pengornis houi</i>.&nbsp; Which is a shame because Luis Chiappe really deserves a bird named after him.&nbsp; So before anyone names a 'Chiappeornis', let's make sure it's a really distinctive specimen, okay?&nbsp; It's also disappointing because O'Connor has an entire section of her thesis lamenting the fact so many enantiornithines are named without adequate diagnoses.&nbsp; But here she's doing that herself (and yes, the contributions credit her with writing the manuscript).&nbsp; I'll also note the paper lacks a measurement table and that Pengornithidae can be paraphyletic in their analysis depending on their assumptions (fig. S1E).&nbsp; Anyway, let's look at the evidence...<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-1VnXredASmo/Voy11XPorDI/AAAAAAAAAv0/Fdxd-dylAS8/s1600/Chiappeavis.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="293" src="http://3.bp.blogspot.com/-1VnXredASmo/Voy11XPorDI/AAAAAAAAAv0/Fdxd-dylAS8/s320/Chiappeavis.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Chiappeavis magnapremaxillo</i> STM 29-11 (after O'Connor et al., 2015).</td></tr></tbody></table><br />Several characters were listed as supposedly diagnostic. The premaxilla was said to have a larger body and convex ventral margin, but the convex element has far too long of a ventral margin to be a premaxilla, so is more likely an incomplete maxilla with the base of the ascending process. The actual right premaxilla is then just the small anterodorsal portion right below the partially preserved left premaxilla, and has no visible ventral margin.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://2.bp.blogspot.com/-O-_z7bSEm4I/Voy8VCbGlEI/AAAAAAAAAwI/CBc5-ZcDm0g/s1600/Chiappeavis%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://2.bp.blogspot.com/-O-_z7bSEm4I/Voy8VCbGlEI/AAAAAAAAAwI/CBc5-ZcDm0g/s320/Chiappeavis%2Bcomp.jpg" width="234" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Left (top to bottom)- skull of <i>Chiappeavis magnapremaxillo</i> as seemingly interpreted by O'Connor et al. (2015b) (top), without interpretation (middle), and as interpreted by me (bottom) (after O'Connor et al., 2015b).&nbsp; Right (top to bottom)- skulls of <i>Parapengornis eurycaudatus</i> holotype IVPP V18687 (top; after Hu et al., 2015), IVPP V18632 (middle; after Hu et al., 2014), and <i>Pengornis houi</i> holotype IVPP V15336 (after O'Connor and Chiappe, 2011).&nbsp; Premaxilla colored red, maxilla colored blue, and nasal colored green.</td></tr></tbody></table><div class="separator" style="clear: both; text-align: center;"></div><br />The posterodorsal premaxillary process was claimed to be longer than other pengornithids, almost reaching the frontals. Yet this bone is highly abraded just posterior to figure 2A's 'l pm' label, so that this posterior portion could easily belong to the right nasal instead. This is bolstered by the fact it's the same width as the left nasal and that the right nasal is otherwise missing. These cranial reinterpretations also make far more sense as they match the morphology and preservation of other pengornithids, which would be expected as the postcrania are nearly identical.<br /><br />The synsacrum has eight vertebrae (as in IVPP V18632; <i>Parapengornis</i>' is only partly preserved), while the authors claim <i>Pengornis</i>' holotype has seven. Yet the description of the latter says only that seven are visible, but that the anterior end is covered, and indeed there could easily be another one beneath the ilium. STM 29-15 (a Jiufotang pengornithid briefly described by O'Connor et al., 2015a) only has seven but is obviously younger based on its unfused sterna.<br /><br />The "median trabeculae" of <i>Chiappeavis </i>are said to have concave lateral margins, but no such structure exists, <strike>so the authors clearly meant the posterolateral processes. Their slight lateral concavity is also seen in <i>Parapengornis</i>' holotype though not in STM 29-15 and perhaps IVPP V18632 (difficult to determine from the low resolution figure of the latter), while <i>Pengornis</i>' holotype doesn't preserve the sternum.</strike> <b>[Edit] O'Connor cordially informed me she meant the posteromedian process, so incorrectly pluralized the term instead of my assumption that she used the wrong positional adjective ("median trabeculae is totally a valid term MORON - they fuse into xiphial region. check Baumel bitch" as per O'Connor pers. comm., 1-16-2016).&nbsp; However, <span><span data-ft="{&quot;tn&quot;:&quot;K&quot;}"><span class="UFICommentBody"><span>concave lateral edges on the posteromedian process are also present in IVPP V18632, absent in STM 29-15, and unknown in both <i>Pengornis </i>and <i>Parapengornis</i>. So this doesn't support <i>Chiappeavis</i>' validity either.</span></span></span></span></b><span style="color: #141823; font-family: Helvetica Neue,Helvetica,Lucida Grande,tahoma,verdana,arial,sans-serif; font-size: 16px; line-height: 21px;"></span><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-n1UzPCSCqoo/VozCTbl8nbI/AAAAAAAAAwk/dNpYMPuROJU/s1600/Parapengornis%2Bsternal%2Bfig.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://4.bp.blogspot.com/-n1UzPCSCqoo/VozCTbl8nbI/AAAAAAAAAwk/dNpYMPuROJU/s320/Parapengornis%2Bsternal%2Bfig.jpg" width="276" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Sternum of <i>Parapengornis eurycaudatus</i> holotype IVPP V18687 showing <strike>concave lateral edge of posterolateral process (right edge of teal line) and</strike> <b>[see discussion above]</b> low posteromedian angle (purple lines) (after Hu et al., 2015).</td></tr></tbody></table><br />The posteromedian angle of the sternum is said to be narrow, but while its 53 degree angle is a bit less than IVPP V18632's (at 68) or STM 29-15's (at 66), <i>Parapengornis</i>' holotype could have an identical angle if complete, and again <i>Pengornis</i>' holotype doesn't preserve the element.<br /><br />Finally, the authors claim the proximal articular surface of the tibia is laterodistally inclined, but this is only true of the left tibia, with the right tibia having a right angle between the surface and the long axis of the bone. Furthermore, <i>Pengornis</i>' holotype and IVPP V18632 both have inclined surfaces, though <i>Parapengornis</i>' holotype lacks them. Given the variation in <i>Chiappeavis</i>' holotype, the variation is likely due to perspective or taphonomy.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-JY0Z_PG6AcA/VozKm8MlFdI/AAAAAAAAAw4/bJUIhUgnDgM/s1600/Pengornis%2Btibiotarsus%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="171" src="http://1.bp.blogspot.com/-JY0Z_PG6AcA/VozKm8MlFdI/AAAAAAAAAw4/bJUIhUgnDgM/s320/Pengornis%2Btibiotarsus%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Tibiotarsi of (left to right; flipped if necessary so that lateral is to the left) both of <i>Pengornis houi</i> holotype IVPP V15336 (after Zhou et al., 2008), right of IVPP V18632 (after Hu et al., 2014; left is in medial view), both of <i>Parapengornis eurycaudatus</i> holotype IVPP V18687 (after Hu et al., 2015), left of STM 29-15 (after O'Connor et al., 2015a; right is fragmented); and <i>Chiappeavis magnapremaxillo</i> holotype STM 29-11 (aftter O'Connor et al., 2015b).&nbsp; Angle of proximal edge indicated in green; note that <i>Chiappeavis</i>' angles differ and that <i>Pengornis</i>' are both ventrolaterally angled.</td></tr></tbody></table><br />Besides the characters listed in the diagnosis, O'Connor et al. note the short anterior cervicals are like the <i>Parapengornis</i> holotype but unlike <i>Pengornis</i>' holotype. As the latter is larger than the others, this may support ontogenetic cervical elongation.<br /><br />They also correctly note the long pygostyle is like <i>Pengornis</i>' holotype, but unlike IVPP V18632 or <i>Parapengornis</i>' holotype.<br /><br /><div style="text-align: left;">Finally, the short metatarsal I is said to be like <i>Pengornis</i>' holotype, and is additionally like IVPP V18632 but unlike <i>Parapengornis</i>' holotype or STM 29-15.&nbsp;</div><div style="text-align: left;"><br /></div><div class="separator" style="clear: both; text-align: left;">Thus the only real difference from <i>Pengornis</i>' holotype is the shorter anterior cervicals, and while there are a few differences from other pengornithid specimens (laterally concave <strike>posterolateral</strike> <b>posteromedian </b>sternal process<strike>es</strike> unlike STM 29-15 <strike>and (?)IVPP V18632</strike>; narrower posterior sternal angle than STM 29-15 and IVPP V18632; long pygostyle unlike STM 29-15, IVPP V18632 and <i>Parapengornis</i>' type; short metatarsal I unlike STM 29-15 and <i>Parapengornis</i>' type), there's no pattern of character distribution that would suggest separate pengornithid species (e.g. <i>Parapengornis </i>shares the cervical length, <strike>posterolateral process concavity</strike> and probably sternal angle, while IVPP V18632 is different in cervical length but shares metatarsal I length).&nbsp; This is illustrated in the following table, showing the distribution of all real differences proposed to diagnose Jiufotang pengornithid taxa (known in more than two specimens) that aren't obviously ontogenetic-&nbsp;</div><div class="separator" style="clear: both; text-align: left;"><br /></div><div class="separator" style="clear: both; text-align: center;"><a href="http://2.bp.blogspot.com/-uOZdph55V2c/Vps7Fjeu1ZI/AAAAAAAAAzU/1YoATUc4-7I/s1600/pengornithid%2Bchar%2Bdist.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" src="http://2.bp.blogspot.com/-uOZdph55V2c/Vps7Fjeu1ZI/AAAAAAAAAzU/1YoATUc4-7I/s1600/pengornithid%2Bchar%2Bdist.jpg" /></a></div><div class="separator" style="clear: both; text-align: center;"></div><div class="separator" style="clear: both; text-align: left;"><br /></div><div style="text-align: left;">B- femoral length (mm) to indicate possible age. </div>C- robust base of maxillary ascending process.<br />D- low interclavicular angle.<br />E- metatarsal I short.<br />F- long anterior cervical vertebrae.<br />G- long pygostyle.<br />H- long posterodorsal lacrimal process.<br />I- laterally concave <strike>posterolateral</strike> <b>posteromedian</b> sternal process.<br />J- narrow posteromedian sternal angle.<br /><br />Note the lack of a pattern unrelated to size.&nbsp; Indeed, all but E, F <b>and I</b> correlate with size, and <b>two of</b> those conflict with <strike>each</strike> <b>the </b>other.&nbsp; It's true that size is not fully correlated with osteological development in this series (e.g. STM 29-15 has unfused sternals unlike the slightly smaller IVPP V18632), but that's true of other taxa as well (e.g. <i>Anchiornis</i>, <i>Archaeopteryx</i>, <i>Shenzhouraptor</i>, <i>Sapeornis</i>, <i>Archaeorhynchus</i>).&nbsp; So it's just like with <i>Archaeopteryx </i>and <i>Microraptor</i>- all specimens show uncorrelated differences, thus either every specimen is its own species or as far as we can tell they're a single species that shows variation.&nbsp; Based on this I conclude all Jiufotang pengornithids should still be still retained in <i>Pengornis houi</i>. <br /><br /><b>References</b>-&nbsp; Zhou, Clarke and Zhang, 2008. Insight into diversity, body size and morphological evolution from the largest Early Cretaceous enantiornithine bird. Journal of Anatomy. 212, 565-577.<br />O'Connor and Chiappe, 2011. A revision of enantiornithine (Aves: Ornithothoraces) skull morphology. Journal of Systematic Palaeontology. 9(1), 135-157.<br />Hu, Zhou and O'Connor, 2014. A subadult specimen of <i>Pengornis </i>and character evolution in Enantiornithes. Vertebrata PalAsiatica. 52(1), 77-97.<br />Hu, O'Connor and Zhou, 2015a. A new species of Pengornithidae (Aves: Enantiornithes) from the Lower Cretaceous of China suggests a specialized scansorial habitat previously unknown in early birds. PLoS ONE. 10(6), e0126791. <br />O'Connor, Wang, Zheng, Hu, Zhang and Zhou, 2015b. An enantiornithine with a fan-shaped tail, and the evolution of the rectricial complex in early birds. Current Biology. <a href="http://dx.doi.org/10.1016/j.cub.2015.11.036" target="_blank">http://dx.doi.org/10.1016/j.cub.2015.11.036 </a><br />O'Connor, Zheng, Sullivan, Chuong, Wang, Li, Wang, Zhang and Zhou, 2015a. Evolution and functional significance of derived sternal ossification patterns in ornithothoracine birds. Journal of Evolutionary Biology. 28(8), 1550-1567.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com9tag:blogger.com,1999:blog-3248412803814730250.post-80581341456554692892015-12-31T21:26:00.001-08:002016-01-04T02:26:29.025-08:00Have a Stremmeia New Year!To finish off the year, here's a taxon of ex-theropod not on anyone's dinosaur lists and which only gets 17 distinct Google results.&nbsp; Perhaps you've heard of the 'Tendaguru <i>Archaeopteryx</i>', a supposed bird carpometacarpus from the Late Jurassic of Tanzania?&nbsp; Well, as Molnar informed me over email, this was actually given a name by Nopcsa in 1930- <i>Stremmeia scabra</i>.&nbsp; Its story follows...<br /><br /><b><i>Stremmeia </i></b>Nopcsa, 1930<br /><b><i>S. scabra</i></b> Nopcsa, 1930<br /><b>Tithonian, Late Jurassic<br />Upper Dinosaur Member of the Tendaguru Formation, Tanzania<br />Holotype</b>- (<strike>HMN</strike> <b>[edit- oh yeah, they changed the abbreviations] MB</b> coll.) ?tibiale (24.5 mm) fused to ?fibulare (27.3)<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-tHm_PfsOGis/VoX3ukMXZ2I/AAAAAAAAAvM/jSBLu4OyRNM/s1600/Stremmeia.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="129" src="http://4.bp.blogspot.com/-tHm_PfsOGis/VoX3ukMXZ2I/AAAAAAAAAvM/jSBLu4OyRNM/s320/Stremmeia.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Stremmeia scabra</i> (HMN coll.) in various views (after Stremme, 1919).</td></tr></tbody></table><br /><b>Comments</b>- This specimen was discovered in 1910 associated with <strike>a <i>Dicraeosaurus </i>skeleton</strike> <b>[edit- Stremme wrote the nonexistent combination '<i>Dicraeosaurus brancai</i>' but specified skeleton S, so meant <i>Brachiosaurus brancai</i>] <i>Giraffatitan</i>'s holotype</b>, initially noted by Janensch (1914) as a bird carpometacarpus probably related to <i>Archaeopteryx</i>, then illustrated and described by Stremme (1919). The latter author could not determine what kind of reptile was represented, or if it was a metacarpus or metatarsus. Lambrecht (1933) noted that Stremme only mentioned <i>Archaeopteryx </i>in the context of differences between its metacarpus and the Tendaguru specimen, so dismissed Parkinson (1930) who stated Stremme considered them relatives. His own opinion was that the carpometacarpus resembled <i>Rhea</i>, so might indicate ratite relationships. Nopcsa (1930) meanwhile had identified the specimen as "carpals" of a pelobatid anuran, citing similarity to the tibiofibulare of <i>Macropelobates </i>(as noted by Stipanicic and Reig, 1957), and named it <i>Stremmeia scabra</i>. Among more recent authors, Hecht (1963) merely said that his reexamination of <i>Stremmeia </i>"showed that these bones are not frog remains" and Estes and Reig (1973) followed his interpretation. There has seemingly been no modern reevaluation of <i>Stremmeia</i>, or discussion of what its identity is if it is not a frog tibiofibulare.<br /><br />Regardless of Hecht's comment, <i>Stremmeia </i>is more similar to e.g. <i>Macropelobates</i>' tibiofibulare than a maniraptoran carpometacarpus in the distinct proximal articular surfaces which form a flat outline instead of a single convex surface, and wide separation of the distal articular surfaces. This latter character is also unlike theropod metatarsals. However, the distal ends are more expanded transversely in anurans, and the distal articular surfaces seem to be simple instead of ginglymoid as in <i>Stremmeia</i>. Further differences from a coelurosaur manus include the large and quadrangular proximal surface of the shorter element (mcIII in most theropods) and sigmoid shape in side view. Whatever <i>Stremmeia </i>turns out to be, it is not theropod. Does anyone have another idea?<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-p60v7t3tIJI/VoX-uDdpyrI/AAAAAAAAAvg/zMEdyJXyEgU/s1600/Stremmeia%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="152" src="http://4.bp.blogspot.com/-p60v7t3tIJI/VoX-uDdpyrI/AAAAAAAAAvg/zMEdyJXyEgU/s320/Stremmeia%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">From left to right- carpometacarpi of <i>Confuciusornis sanctus</i> (Bonn specimen; after Goernemann, 1999) and <i>Anchiornis huxleyi</i> (holotype IVPP V14378; after Xu et al., 2008); metacarpals II and III of <i>Archaeopteryx lithographica</i> (Eichstatt specimen JM SoS 2257; after Wellnhofer, 1974); <i>Stremmeia scabra</i> (holotype HMN coll.; after Stremme, 1919); and tibiofibulare of <i>Macropelobates osborni</i> (holotype AMNH 6252; after Noble 1924).</td></tr></tbody></table><br /><b>References</b>- Janensch, 1914. Ubersicht uber die Wirbeltierfauna der Tendaguru-Schichten. Archiv fur Biontologie. 3, 81-110.<br />Stremme, 1919. Uber die durch Bandverknocherung hervorgerufene proximale Verschmelzung zweier Mittelhand - oder Mittelfussknochen eines Reptils. Wissenschaftliche Ergebnisse der Tendaguru-Expedition. Archiv fur Biontologie. 4, 143-144.<br />Noble, 1924. A new spadefoot toad from the Oligocene of Mongolia with a summary of the evolution of the Pelobatidae. American Museum Novitates. 132, 15 pp.<br />Nopcsa, 1930. Notes on Stegocephalia and Amphibia. Proceedings of the Zoological Society of London. 1930, 979-995. <br />Parkinson, 1930. The dinosaur in East Africa: An account of the giant reptile beds of Tendaguru, Tanganyika territory. H.F. &amp; G. Witherby. 192 pp.<br />Lambrecht, 1933. Handbuch der Palaeornithologie. Gebruder Borntraeger. 1024 pp. <br />Stipanicic and Reig, 1957. El "Complejo Porfírico de la Patagonia extraandina" y su fauna de anuros. Acta Geologica Lilloana. 1, 185-297.<br />Hecht, 1963. A reevaluation of the early history of the frogs. Part II. Systematic Zoology. 12(1), 20-35.<br />Estes and Reig, 1973. The early fossil record of frogs: A review of the evidence. In Vial (ed.). Evolutionary biology of the anurans: Contemporary research on major problems. University of Missouri Press. 11-63.<br />Wellnhofer, 1974. Das fünfte Skelettexemplar von <i>Archaeopteryx</i>. Palaeontographica. 147, 169-216. <br />Goernemann, 1999. Osteologie eines Exemplars von <i>Confuciusornis</i> aus der unteren Kreide von West-Liaoning, China. Archaeopteryx. 17, 41-54.<br />Xu, Zhao, Norell, Sullivan, Hone, Erickson, Wang, Han and Guo, 2008. A new feathered maniraptoran dinosaur fossil that fills a morphological gap in avian origin. Chinese Science Bulletin. 54(3), 430-435.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-27832491488374463642015-10-19T01:29:00.000-07:002015-10-19T13:28:46.305-07:00SVP 2015 Day 4The final day.&nbsp; Let's see what happened...<br /><br />In another ornithischian abstract, Borinder et al. redescribe the hadrosaur <i>Tanius</i>.&nbsp; As the authors state, it's been 86 years since the genus was described.&nbsp; The only known specimen is immature and has a flexor canal on the femur.&nbsp; Unfortunately, it emerges in a big polytomy with other taxa just outside Hadrosauridae.&nbsp; I do wonder if excluding some of these nine taxa a posteriori might show some resolution between the remaining ones and <i>Tanius</i>.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-54zSF1s9hQk/ViScTBMaE9I/AAAAAAAAAuc/4elUgcBD0DM/s1600/Tanius%2Bskull.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="188" src="http://4.bp.blogspot.com/-54zSF1s9hQk/ViScTBMaE9I/AAAAAAAAAuc/4elUgcBD0DM/s320/Tanius%2Bskull.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Cranial elements of <i>Tanius sinensis</i> holotype (after Wiman, 1929).</td></tr></tbody></table><br />Cuesta et al. argue that the ulnar bumps of <i>Concavenator </i>are homologous to the secondary remix attachment points of birds.&nbsp; This was disputed separately by Naish and <a href="http://theropoddatabase.blogspot.com/2010/09/concavenator-part-ii-becklespinax.html" target="_blank">myself shortly after the genus was described</a>.&nbsp; I still don't see how the authors interpret the bumps as posterolateral instead of anterolateral.&nbsp; I argued the structure was an intermuscular line between the flexor ulnaris and the extensor carpi radialis brevis (sensu Meers, 2003), or flexor digitorum profundus and extensor carpi ulnaris (sensu Gishlick, 2002).&nbsp; Cuesta et al. state that (besides the triceps brachii which we both agree is back on the olecranon) they reconstructed the anconeus and abductor polices longus and that the bumps are not located between them.&nbsp; The abductor polices longus (pronator quadratus in Meers) is a more proximodorsally located scar I agree has nothing to do with the bumps.&nbsp; I think anconeus is another term for the extensor carpi ulnaris, based on Gishlick's figure of <i>Corvus </i>and Hudson and Lanzillotti's (1964) description.&nbsp; So with the caveats that there seems to be no consensus for muscle names or homology between crocodylians and birds, I don't think Cuesta et al. had the same muscles in mind as I did.<br /><br />Fortner reports "parts of an associated postcranial skeleton of a small theropod dinosaur recently collected from the uppermost Aguja Formation."&nbsp; This is another frustrating example of the information-bereft abstract, with the first twelve lines devoted to introduction and background knowledge.&nbsp; The only other bit of information is- "The specimen exhibits some unique features, but is compatible with identification as either Troodontidae or Dromaeosauridae."&nbsp; This is very intriguing, as while both eudromaeosaur and troodontine teeth are known from the Aguja, we have <i>Richardoestesia </i>and <i>Paronychodon</i> from there too.&nbsp; As I think these are microraptorine and basal troodontid respectively, this specimen could fit the bill.&nbsp; Did anyone get more details?<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-EtKFl7_qtIk/ViSdYf109TI/AAAAAAAAAuk/ugsdlju1aOI/s1600/Paronychodon%2Bdentary.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://3.bp.blogspot.com/-EtKFl7_qtIk/ViSdYf109TI/AAAAAAAAAuk/ugsdlju1aOI/s320/Paronychodon%2Bdentary.jpg" width="291" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">The only published non-dental remains attributed to <i>Paronychodon</i>- a partial dentary IPFUB GUI D 1 in A. medial, B. dorsal, and C. lateral views (after Zinke and Rauhut, 1994).&nbsp; It does differ from later <i>Paronychodon </i>in having distal and sometimes mesial serrations.</td></tr></tbody></table><br />Harding et al. have the non-dinosaurian abstract I think could be most important to dinosaur studies.&nbsp; They examined all "published characters for discriminating <i>Sceloporus </i>from the related iguanian lizards <i>Uta </i>and <i>Urosaurus</i>, and for discriminating among species of <i>Sceloporus</i>" for 14 species, "11 of [which] had more than ten individual skeletal specimens, and for four of them sample sizes exceeded 50 specimens."&nbsp; Perhaps astonishingly "almost all characters published in the literature to identify fossil specimens have no power to discriminate reliably between <i>Uta</i>, <i>Urosaurus</i>, and <i>Sceloporus</i>, nor between species of <i>Sceloporus </i>we examined."&nbsp; In our field where most species are only known from single specimens, and most known from multiple specimens don't have more than one or two described in detail, what does this mean for our autapomorphy lists?<br /><br />Holtz et al. report more information on an <i>Anzu </i>specimen (anyone know the collection number?) they announced last year.&nbsp; Interestingly, only distal tarsal IV is fused to the unfused metatarsus, and "a pair of pronounced cruciate ridges on the plantar surface of metatarsal III" are present as in <i>Elmisaurus </i>but don't extend as far proximally.&nbsp; This would make it intermediate in pedal morphology between <i>Chirostenotes </i>and <i>Elmisaurus</i>.&nbsp; <br /><br />Maddin et al. present a hypothesis based on the development of skull roof bones in mice and chickens.&nbsp; In mice (and axolotls) the suture between the frontal and parietal corresponds to the boundary between the neural crest and mesoderm in the embryo.&nbsp; In chickens, the latter embryonic boundary is within the frontal itself.&nbsp; Thus the authors suggest the avian 'frontal' is actually a fused frontal and parietal, while the avian 'parietal' is actually a postparietal.&nbsp; They say that hypothesis "is also supported phylogenetically where data from the fossil record reveal separate frontal, parietal, and postparietal bones are present in all stem lineages of extant taxa, including that of birds (e.g., the stem archosaur <i>Euparkeria</i>)."&nbsp; The problem I see is that the postparietal in basal archosauriforms is a tiny wedge between the parietals and supraoccipital, while the parietals have the same topological relationships and morphology in these basal archosauriforms that they do in basal dinosaurs.&nbsp; It seems more likely to me that this is a case like manual homology where the developmental process itself evolves, so that somewhere on the sauropsid line, the neural crest-mesoderm boundary moved into the frontal.&nbsp; Also interesting would be to know what the state in lepidosaurs and crocodylians is.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://2.bp.blogspot.com/-oNTcdeG4KV4/ViSfzzGUd3I/AAAAAAAAAuw/emyAjq6dvnA/s1600/Maddin%2Bet%2Bal%2Bskull%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="123" src="http://2.bp.blogspot.com/-oNTcdeG4KV4/ViSfzzGUd3I/AAAAAAAAAuw/emyAjq6dvnA/s320/Maddin%2Bet%2Bal%2Bskull%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Comparison of skull roof in dorsal view of- left, <i>Erythrosuchus africanus</i> (after Gow, 2003); center, <i>Ornithosuchus longidens</i> (after Walker, 1964), and right,<i> Herrerasaurus ischigualastensis</i> (after Sereno and Novas, 1993).&nbsp; Parietal is blue, postparietal is red.&nbsp; Maddin et al. suggest the red at left is homologous to the blue at right.</td></tr></tbody></table><br />Mannion et al. redescribe the famous 'French <i>Bothriospondylus</i>' (MNHN coll.), presumably based on Moine's (1999) thesis.&nbsp; It's great that we're getting all of the historical <i>Bothriospondylus </i>material redescribed in the last decade.<br /><br />McFeeters et al. do the important job of reevaluating <i>Struthiomimus </i>specimens.&nbsp; Little known to most, the holotype is extremely fragmentary, with most information coming from Osborn's AMNH 5339, Nicholls and Russell's UCMZ 1980.1 and the new RTMP 90.26.1. The authors find "a relatively small partial skeleton from the lower Dinosaur Park Formation" (which based on listed elements must be ROM 1970) to belong to a new taxon of ornithomimid based on several autapomorphies.&nbsp; This specimen forms the basis of the dorsal snout in Russel's (1972) and Paul's (1988) cranial reconstructions, making those composites.&nbsp; Another specimen shares some pelvic characters with <i>Qiupalong</i>, perhaps relating to a Dinosaur Park astragalus reported by McFeeters et al. in last year's SVP abstract.&nbsp; <br /><br />Nesbitt et al. seem set to provide a detailed description of <i>Asilisaurus</i>, after the original tabloid announcement.&nbsp; Interestingly, <i>Agnosphitys </i>is said to a basal silesaurid as well, which is the fourth proposed identification for the genus.&nbsp; Add this to Agnolin's (2015) Jornadas Argentinas abstract proposing <i>Pisanosaurus </i>belongs to the clade, and its membership is expanding markedly.<br /><br />We also get yet another description of <i>Nothronychus</i>' braincase.&nbsp; Far be it for me to complain about having too many descriptions of a taxon, but this one specimen of one taxon was described in 2005, 2012 and 2013, and now we'll probably get a fourth portion next year or so.&nbsp; At least this new abstract is about the previously unrecognized anterior portion, but I'd rather have that time and effort go towards... say "Zunityrannus" or describing the Bayan Shiree therizinosaur postcrania.&nbsp; Smith et al. report that "there is a supraorbital evagination in this specimen that is currently interpreted as accommodating a well-developed nasal gland in the frontal.&nbsp; This development has been observed in some other archosaurs, especially marine birds, where it is associated with salt excretion and is consistent with a beach or other evaporitic paleoenvironmental interpretation."&nbsp; So let's all start our All Yesterdays style marine therizinosaur pics.<br /><br />Finally for this year, Sullivan et al. report a 'sphenosuchian' specimen sister to <i>Junggarsuchus</i>.&nbsp; This has the interesting mix of cursorial features with a webbed hand and distal tail sheathed with armor.&nbsp; <br /><br /><b>References</b>- Wiman, 1929. Die Kreide-Dinosaurier aus Shantung. Palaeontologia Sinica (series C). 6, 1-67.<br /><br />Hudson and Lanzillotti, 1964. Muscles of the pectoral limb in galliform birds. The American Midland Naturalist. 71(1), 1-113.<br /><br />Walker, 1964. Triassic reptiles from the Elgin area: <i>Ornithosuchus </i>and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London, seies B. 248(744), 53-134.<br /><br />Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada. Canadian Journal of Earth Sciences. 9, 375-402.<br /><br />Paul, 1988. Predatory Dinosaurs of the World. Simon &amp; Schuster: New York. 464 pp.&nbsp; <br /><br />Sereno and Novas, 1993. The skull and neck of the basal theropod <i>Herrerasaurus ischigualastensis</i>. Journal of Vertebrate Paleontology. 13, 451-476.<br /><br />Zinke and Rauhut, 1994. Small theropods (Dinosauria, Saurischia) from the Upper Jurassic and Lower Cretaceous of the Iberian Peninsula. <span class="st">Berliner Geowissenschaftliche Abhandlungen</span>. E 13, 163-177.<br /><br />Moine, 1999. Datation, condition de depot et position phylogenetique de '<i>Bothriospondylus madagascariensis</i>' (Damparis,<br />Jura, France). MS thesis, Memoires de Maıtrise Magistere Sciences de la Terre ENS-Lyon.<br /><br />Gishlick, 2002. The functional morphology of the forelimb of <i>Deinonychus antirrhopus</i> and its importance for the origin of avian flight. PhD thesis, Yale University. 142 pp.<br /><br />Gower, 2003. Osteology of the early archosaurian reptile <i>Erythrosuchus africanus</i> Broom. Annals of the South African Museum. 110(1), 1-88.<br /><br />Meers, 2003. Crocodylian forelimb musculature and its relevance to Archosauria. The Anatomical Record. Part A, 274A, 891-916.<br /><br />Agnolin, 2015.&nbsp; Nuevas observaciones sobre <i>Pisanosaurus mertii</i> Casamiquela, 1967 (Dinosauriformes) y sus implicancias taxonomicas. XXIX Jornadas Argentinas de Paleontologia de Vertebrados. Libros de Resumenes. 13-14.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com17tag:blogger.com,1999:blog-3248412803814730250.post-35841768897412608172015-10-16T14:23:00.000-07:002015-10-16T14:23:53.653-07:00SVP 2015 Day 3Time for day three.&nbsp; If I were in Dallas, I'd be waking up early for Technical Session X.<br /><br />Carrano and Choiniere reexamine <i>Ceratosaurus</i>' forelimb based on the holotype.&nbsp; The whole thing could use redescription since Gilmore's last one 95 years ago.&nbsp; While they state "These new data are consistent with the placement of <i>Ceratosaurus </i>as close to (or within) Abelisauroidea", an abelisauroid <i>Ceratosaurus </i>is impossible as Ceratosauroidea predates Abelisauroidea, without even getting into phylogenetic nomenclature.&nbsp; <br /><br />Hey, we get more <i>Dromiceiomimus </i>feathers.&nbsp; van der Reest et al. report on a new specimen preserving feathers on the proximal thigh and dorsal tail, but not ventral tail or distal hindlimb.&nbsp; The feathers are also branched, moving this trait to the base of Maniraptoriformes.&nbsp; As the feathers have rachis but lack barbules, it seems Prum's Stage IIIa was correct.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://2.bp.blogspot.com/-BlSMPmC62U0/ViFFHtXE6jI/AAAAAAAAAt0/jcCEf2RUV18/s1600/Segnosaurus%2B100%2B83%2Bradius%2Bulna.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="240" src="http://2.bp.blogspot.com/-BlSMPmC62U0/ViFFHtXE6jI/AAAAAAAAAt0/jcCEf2RUV18/s320/Segnosaurus%2B100%2B83%2Bradius%2Bulna.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Segnosaurus galbinensis</i> radius and ulna (paratype IGM 100/83), courtesy of Zanno.</td></tr></tbody></table><br />Kobayashi et al. describe a new therizinosaur specimen from the Bayan Shiree Formation, where <i>Segnosaurus</i>, <i>Erlikosaurus</i> and <i>Enigmosaurus </i>come from.&nbsp; While it's notable for having a reduced metacarpal III (and thus perhaps reduced third manual digit), none of the three named genera from that formation preserve enough manual material to evaluate their condition.&nbsp; The new specimen is potentially comparable to <i>Enigmosaurus </i>and <i>Segnosaurus </i>in that all preserve the radius and ulna.<br /><br />Funston and Currie report on their fairly complete Horseshoe Canyon caenagnathid.&nbsp; Supposedly having cervicals "distinct from <i>Epichirostenotes</i>", an "articular ridge is intermediate in size and form between <i>Caenagnathus collinsi</i>" and <i>sternbergi</i>, and autapomorphic manual proportions, it should be useful for resolving caenagnathid taxonomy.&nbsp; In addition to the expected anatomical details, ulnar "feather scars" are said to be present.<br /><br />Lu et al. discuss the billionth new oviraptorid from the Nanxiong Formation.&nbsp; This one's based on at least a skull and is similar to <i>Khaan</i>.&nbsp; Some of these things have got to be synonymous <br /><br /><i>Saurornitholestes </i>has been the ubiquitous but rarely described Late Cretaceous American dromaeosaurid.&nbsp; We've had RTMP 88.121.39 and MOR 660 known since 1988 and 1990 respectively, but both are only mentioned in passing in papers.&nbsp; Now we have a new almost complete specimen discovered in 2014 (did anyone who saw the talk catch the specimen number?) that includes a skull.&nbsp; Is the <i>Saurornitholestes </i>osteology finally at hand?<br /><br />Perhaps the most unusual theropod abstract is from Sorkin, describing a phylogeny for tetanurines.&nbsp; I'm not sure if it's based on a quantitative analysis or just the noted 'key characters', but it's not similar to the current consensus.&nbsp; From the abstract, his topology and taxonomy seems to be-<br /><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">|--Ceratosauria<br />`--Tetanurae<br />&nbsp;&nbsp; |--Spinosauridae</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp;&nbsp; |--Carcharodontosauridae</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp;&nbsp; `--+--<i>Sinosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; |--Piatnitzkysauridae</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; |--Metriacanthosauridae</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; |--<i>Neovenator </i></span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; |--Megaraptora</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; `--Avetheropoda</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |--Megalosauridae</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--Euavetheropoda</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--<i>Monolophosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--Acrocanthosauridae (incl. <i>Eocarcharia</i>)</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--Allosauridae</span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--Coelurosauria </span><br /><br />Brings you back to the early 90s, doesn't it?&nbsp; Note the new clade name Euavetheropoda, said to be distinguished by a robust dorsal quadratojugal process.&nbsp; In any case, the accepted phylogenetic definition for Avetheropoda would put it at the Allosauridae+Coelurosauria node, making Euavetheropoda unintuitively more inclusive.&nbsp; <br /><br />Wills describes sixteen teeth from the Bathonian Forest Marble Formation of England.&nbsp; When analyzed, these plot with dromaeosaurids.&nbsp; He states "This pushes back the origin of dromaeosaurids from the Kimmeridgian to the Bathonian", but Metcalf and Walker described teeth from the Bathonian Chipping Norton Formation of England as dromaseosaur-like back in 1994.&nbsp; Maybe these new ones are more securely identified.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-eWBapuDUJQw/ViFl3nHE5eI/AAAAAAAAAuI/RKGm6YumlI8/s1600/Chipping%2BNorton%2Bdrom%2Btooth.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="294" src="http://3.bp.blogspot.com/-eWBapuDUJQw/ViFl3nHE5eI/AAAAAAAAAuI/RKGm6YumlI8/s320/Chipping%2BNorton%2Bdrom%2Btooth.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Dromaeosaurid-like tooth (GLRCM G.51422) from the Bathonian Chipping Norton Formation in lingual (A), labial (B) and basal (C) views (after Metcalf and Walker, 1994).</td></tr></tbody></table>Just one more day to go, with quite a few interesting abstracts tomorrow.<br /><br /><b>Reference</b>- Metcalf and Walker, 1994. A new Bathonian microvertebrate locality in the English Midlands. In Fraser and Sues (eds.). In the Shadow of the Dinosaurs- Mesozoic Small Tetrapods. Cambridge University Press. 322-332.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-12772089517413559922015-10-15T22:45:00.003-07:002015-10-15T22:45:59.561-07:00SVP 2015 Day 2And we're on to day two.&nbsp; Something I noticed about this year's and last year's abstracts is just how many of them are already published in official format by the time the meeting happens (yet another reason the embargo is silly).&nbsp; I gather one of the big issues facing SVP is how many talks and papers there are, leading to greater expense, more parallel sessions, etc..&nbsp; If the dinosaur abstracts are any indication, you could cut out a third of them by excluding those that will be published by September. <br /><br />Pritchard created a new matrix to test the relationships of Sauria, but alas this is one of those abstracts that doesn't actually contain much information.&nbsp; The most it says is that Protorosauria is para/polyphyletic, which everyone agrees with by now.&nbsp; I would ask that if your SVP abstract is based on a phylogenetic analysis, please devote at least a couple sentences to describing the topology you found.&nbsp; Otherwise it's just a tease and I learn nothing.<br /><br />This was a great SVP for ornithischians.&nbsp; We've had Arbour revise ankylosaurids, and now Burns is doing the same for Campanian-Maastrichtian North American nodosaurids. He finds <i>Denversaurus </i>is a valid taxon, sister to <i>Panoplosaurus</i>.&nbsp; So that's another genus from your 1980s dino encyclopedias to dust off.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-EWMUOvB6u3U/ViBe3swI7aI/AAAAAAAAAtM/W_6iAluXq0Y/s1600/Denver.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="240" src="http://1.bp.blogspot.com/-EWMUOvB6u3U/ViBe3swI7aI/AAAAAAAAAtM/W_6iAluXq0Y/s320/Denver.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Denversaurus</i>.&nbsp; What?&nbsp; That's not right?...</td></tr></tbody></table><br />Continuing the ornithischian train, Barta and Norell report on new specimens of <i>Haya</i>.&nbsp; The interesting thing here is that they performed two analyses- one with each specimen coded as a separate OTU, and the other with one <i>Haya </i>OTU that was coded as polymorphic when specimens differed.&nbsp; In the first, <i>Haya </i>emerged as a basal thescelosaurid, but in the second it was a basal neornithischian.&nbsp; This is presumably because PAUP/TNT finds it most parsimonious to choose a mix of states for the polymorphic characters that isn't found in any actual specimen.&nbsp; It's concerning because being a lumper myself, I code e.g. <i>Microraptor </i>and <i>Archaeopteryx </i>as single OTUs.&nbsp; Is that affecting their relationships in my analyses?&nbsp; <br /><br />Shelley et al.'s abstract is an example of two things I like.&nbsp; First, figuring out where all of those extinct mammal groups go using a molecular scaffold for the topology.&nbsp; Second, actually describing the results of the study- "Our phylogenetic analysis places "triisodontids" as a basal member of Euungulata within Laurasiatheria. "Triisodontidae" forms a paraphyletic stem of Mesonychia with <i>Oxyclaenus </i>most closely related to a monophyletic Mesonychia.&nbsp; "Triisodontids" plus Mesonychia are closely related to a clade comprised of the arctocyonids <i>Mimotricentes</i>, <i>Deuterogonodon </i>and <i>Chriacus</i>."&nbsp; Ahhh, actual information...<br /><br />Besides the usual morass of Yixian and Jiufotang birds (including <i>Parapengornis</i>, <a href="http://theropoddatabase.com/Ornithothoraces.htm#Pengornishoui" target="_blank">which I think is just <i>Pengornis</i></a>), we get another specimen from the lower member of the Huajiying Formation.&nbsp; This earlier horizon has otherwise only yielded <i>Confuciusornis zhengi</i>, <i>Protopteryx</i>, <i>Eopengornis </i>and <i>Archaeornithura</i>.&nbsp; Hu et al.'s new enantiornithine is said to have a <i>Liaoningornis</i>-like sternum, which could cement the affinities of that genus.&nbsp; <br /><br />The Norman-Barrett team's on the basal ornithischian case again, this time with Baron et al.'s redescription of <i>Lesothosaurus</i> postcrania.&nbsp; This is needed, as Sereno (1991) mostly described the skull and thus we've had to depend on Thulborn's work from 43 years ago.&nbsp; They find <i>Stormbergia </i>to be based on older individuals of <i>Lesothosaurus</i>, which as a lumper, does not surprise me.&nbsp; The genus emerges as a basal neornithischian.&nbsp; This should be a good paper once it's published.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-pZXzg8giQqQ/ViCKPCNN7pI/AAAAAAAAAtg/ToF9xx2RjGs/s1600/Morosaurus%2Bagilis.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="120" src="http://1.bp.blogspot.com/-pZXzg8giQqQ/ViCKPCNN7pI/AAAAAAAAAtg/ToF9xx2RjGs/s320/Morosaurus%2Bagilis.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>"Morosaurus" agilis</i> (USNM 5384) posterior skull and anterior cervicals in left lateral view (after Gilmore, 1907).</td></tr></tbody></table><br />Finally, Whitlock and Wilson redescribe the hitherto enigmatic <i>"Morosaurus" agilis</i>.&nbsp; Based on a braincase and anterior cervicals, it turns out to be a diplodocid.&nbsp; While apparently not <i>Apatosaurus </i>(in which the abstract seems to include <i>Brontosaurus</i>) or <i>Galeamopus</i>, the newly exploded Morrison Diplodocidae leaves open numerous possible identifications- <i>Supersaurus</i>, <i>Amphicoelias</i>, <i>Kaatedocus</i>, <i>Barosaurus</i>, <i>Diplodocus</i>...&nbsp; I'm not sure I believe its affinities can't be narrowed down further.&nbsp; For instance, Lovelace et al. (2007) stated small cervical pleurocoels were diagnostic for <i>Supersaurus</i>, and <i>agilis </i>has large pleurocoels.&nbsp; Tschopp and Mateus (2013) proposed numerous characters to distinguish <i>Kaatedocus </i>from other diplodocids, including a postorbitally restricted squamosal that <i>agilis </i>seems to have, and a postparietal foramen <i>agilis</i> seems to lack.&nbsp; Maybe published characters have issues that I'm not aware of as a theropod worker, or maybe Gilmore's description is misleading, but I find hard to believe that something as complex as a braincase and posterior skull can't be distinguished between <i>Kaatedocus </i>and <i>Diplodocus</i> (even if <i>Amphicoelias </i>and <i>Barosaurus </i>can't be compared).<br /><br />Join me again tomorrow, when we open with those sweet, sweet theropod abstracts...<br /><br /><br /><b>References</b>- Gilmore, 1907. The type of the Jurassic reptile <i>Morosaurus agili</i>s redescribed, with a note on <i>Camptosaurus</i>. Proceedings of the United States National Museum. 32(1519), 151-165.<br /><br />Sereno, 1991. <i>Lesothosaurus</i>, "fabrosaurids," and the early evolution of Ornithischia. Journal of Vertebrate Paleontology. 11(2), 168-197.<br /><br />Lovelace, Hartman and Wahl, 2007. Morphology of a specimen of <i>Supersaurus </i>(Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional Rio de Janeiro. 65, 527-544.<br /><br />Tschopp and Mateus, 2013. The skull and neck of a new flagellicaudatan sauropod from the Morrison Formation and its implication for the evolution and ontogeny of diplodocid dinosaurs. Journal of Systematic Palaeontology. 11, 853-888.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com4tag:blogger.com,1999:blog-3248412803814730250.post-86208995553717670312015-10-14T20:03:00.003-07:002015-10-14T20:03:49.582-07:00SVP 2015 Day 1While again too poor to attend SVP, I thought I would provide my thoughts on the abstracts that interest me like I did last year.&nbsp; The abstracts book can be download free at <a href="http://vertpaleo.org/Annual-Meeting/Home.aspx" target="_blank">this link</a>.&nbsp; When adding abstracts to the Database, I noticed an issue that's been constant throughout the years- the titles are in UPPERCASE.&nbsp; This makes copying them useless, which means everyone citing abstracts has to rewrite them.&nbsp; This can only lead to more typos and serves no obvious purpose since the titles are already bolded to distinguish them from the rest of the text.&nbsp; Does anyone else agree they should have normal capitalization?<br /><br />This year there are no less than three abstracts marked as WITHDRAWN.&nbsp; One is by Egberts and concerns wearing gloves when handling specimens to prevent skin oils damaging the fossils.&nbsp; Another is by Spindler, about a supposed Carboniferous therapsid specimen.&nbsp; The third by Parsons and Parsons involves aerial forelimb motion in <i>Bambiraptor </i>and <i>Deinonychus</i>.&nbsp; I wonder why these three abstracts didn't make it.<br /><br />Wilson reports that of three histologically sampled <i>Pteranodon </i>specimens, the largest and smallest are rather mature, while the medium-sized one is immature.&nbsp; While she interprets this to mean "there may be a large amount of adult body size variation in <i>Pteranodon</i>" or possibly that "the smallest specimen sampled is from a large <i>Nyctosaurus </i>specimen", Peters' idea of taxonomic instead of ontogenetic variation in <i>Pteranodon </i>seems viable too.&nbsp; Is this a case of Peters being right?<br /><br />Andres (and posthumously Langston) signal the beginning to the end of the deplorable situation pterosaur workers have been in where <i>Quetzalcoatlus</i>' holotype has been inaccessible due to the <a href="http://pterosaur-net.blogspot.com/2010/05/embarrassing-questions-on.html" target="_blank">TMM embargoing it for decades</a>.&nbsp; All it took was the eventual death of the person monographing it.&nbsp; :|&nbsp; Now if we can just get <i>Pelecanimimus </i>out of the same rut, so that those who have the thesis describing it will distribute it and allow others to photograph the material described over two decades ago...<br /><br /><div class="separator" style="clear: both; text-align: center;"><a href="http://4.bp.blogspot.com/-G7evswVrjGk/Vh8PMCi_C0I/AAAAAAAAAs8/sEyEHmagAYY/s1600/Bellusaurus%2Bfrontal%2Bcomp.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" height="165" src="http://4.bp.blogspot.com/-G7evswVrjGk/Vh8PMCi_C0I/AAAAAAAAAs8/sEyEHmagAYY/s320/Bellusaurus%2Bfrontal%2Bcomp.jpg" width="320" /></a></div>Frontals of <i>Bellusaurus sui</i> referred specimen IVPP V17768.7 (left; after Mo, 2013), <i>Europasaurus holgeri</i> referred specimen DFMMh/FV 162 (center; after Marpmann et al., 2015), and <i>Camarasaurus lentus</i> referred specimen CM 11338 (right; after Gilmore, 1925) showing two characters reported by Moore et al. as shared between the former two genera- elongate frontal and deep orbital concavity in frontal.&nbsp; <br /><br /><br />Moore et al. report on new <i>Bellusaurus </i>cranial elements, juvenile like the previously known material.&nbsp; Interestingly, these support a macronarian position, and close relationship with <i>Europasaurus</i>.&nbsp; No update on the oft-hypothesized synonymy with <i>Klamelisaurus</i>, as that genus only preserves teeth and postcrania.<br /><br />Finally, we have the abstract supposedly authored by Chinzorig, Kobayashi, Tsogtbaatar, Mahito, Rinchen and Shigeru...?&nbsp; Someone messed up somewhere, because of course the actual authorship using surnames should be Tsogtbaatar, Kobayashi, Tsogtbaatar, Watabe, Barsbold and Suzuki.&nbsp; This one's going to be tedious to find/cite in the future.&nbsp; The find itself is a diagnostic ornithomimid tarsus and pes from the Djadokhta Formation of Mongolia, unfortunately not comparable to the previously described cranial and vertebral material (IGM 100/987 and 100/1245) from that formation.<br /><br />Join me tomorrow for Day 2 talks and posters...<br /><br /><b>References</b>- Gilmore, 1925. A nearly complete articulated skeleton of <i>Camarasaurus</i>, a saurischian dinosaur from the Dinosaur National Monument. Memoirs of the Carnegie Museum. 10, 347-384.<br /><br />Mo, 2013. <i>Bellusaurus sui</i>. Topics in Chinese Dinosaur Paleontology. Henan Science and Technology Press. 155 pp.<br /><br />Marpmann, Carballido, Sander and Knötschke, 2015. Cranial anatomy of the Late Jurassic dwarf sauropod <i>Europasaurus holgeri</i> (Dinosauria, Camarasauromorpha): Ontogenetic changes and size dimorphism. Journal of Systematic Palaeontology. 13(3), 221-263.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-48232148119234177242015-09-17T19:36:00.002-07:002015-09-17T19:36:52.794-07:00Scaphonyx, Hyperodapedon minor and a site updateThe final example of four-ex-saurischians is <i>Scaphonyx fischeri</i>.&nbsp; A big thanks to Mike Benton for helping me with <i>Hyperodapedon minor</i>.&nbsp; Today <a href="http://theropoddatabase.com/Updates.htm" target="_blank">The Theropod Database was also updated</a>, and the next update in October will feature all the new SVP 2015 information.<br /><br /><i><b>Hyperodapedon </b></i>Huxley vide Murchison, 1858<br />= <i>Stenometopon </i>Boulenger, 1904<br />= <i>Scaphonyx </i>Woodward, 1907<br />= <i>Cephalastron </i>Huene, 1926<br />= <i>Cephalonia </i>Huene, 1926<br />= <i>Cephalostronius </i>Huene, 1926<br />= <i>Scaphonychimus </i>Huene, 1926<br />= <i>Macrocephalosaurus </i>Tupi-Caldas, 1933 <br />= <i>Paradapedon </i>Huene, 1938<br />= <i>Supradapedon </i>Chatterjee, 1980<br /><b>Definition</b>- (<i>Hyperodapedon gordoni</i><!-----> &lt; - <i>Teyumbaita sulcognathus</i>) (Langer and Schultz, 2000)<br /><b>References</b>- Murchison, 1858. On the sandstones of Morayshire (Elgin, &amp;c.) containing reptilian remains; and on their relations to the Old Red Sandstone of that country. Quarterly Journal of the Geological Society of London. 15, 419-439.<br />Boulenger, 1904. On reptilian remains from the Triass of Elgin. Philosophical Transactions of the Royal Society of London B. 196, 175-189. <br />Woodward, 1907. On some fossil reptilian bones from the state of Rio Grande do Sul. Revista do Museu Paulista. 7, 46-57. <br />Huene, 1926. Gondwana-Reptilien in Südamerika. Palaeontologia Hungarica. 2, 1-108. <br />Tupi-Caldas, 1933. Contribuição ao estudo do fossil da Alemoa, Município de Santa Maria, Rio Grande do Sul. In Tupi-Caldas (ed.). Curso Geral de Mineralogia e Geologia, aplicada ao Brasil. Edições da Livraria do Globo. 333-339. <br />Huene, 1938. <i>Stenaulorhynchus</i>, ein Rhynchosauridae der ostafrikanischen Obertrias. Nova Acta Leopoldina. 1938, 83-121. <br />Chatterjee, 1980. The evolution of rhynchosaurs. Memoires de la Societe Geologique de France, Nouvelle Serie. 139, 57-65. <br />Langer, 1996. Rincossauros sul-brasileiros: Historico e filogenia. Masters thesis, Universidade Federal do Rio Grande do Sul. 361 pp.<br /><br />&nbsp;<i><b>H. fischeri</b></i> (Woodward, 1907) Whatley, 2005<br />= "Scaphonyx fischeri" White, 1906<br />= <i>Scaphonyx fischeri</i> Woodward, 1907<br /><b>Carnian, Late Triassic<br />Alemoa Member of the Santa Maria Formation, Brazil<br />Holotype</b>- (BM R-5033) two cervical centra, dorsal centrum, central fragment, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, pedal ungual I <br /><br /><b>Comments</b>- The holotype was discovered in 1902, and initially announced by Woodward in 1903 before being described and named by him in 1907. White (1906) first published the name in a note in Science, but did not provide a description or definition (ICZN Article 12.1), making the name a nomen nudum. Woodward (1907) identified <i>Scaphonyx </i>as a <i>Euskelosaurus</i>-like dinosaur based on several characters. First, the dorsal centrum lacks a parapophysis, supposedly unlike 'anomodonts' (under which he included pareiasaurs, procolophonids and therapsids), but rhynchosaurs (which Woodward classified as rhynchocephalians) possess the same state as <i>Scaphonyx</i>. Second, the cervical supposedly resembled <i>Euskelosaurus</i>, but this was based on a specimen (BMNH R2791) now referred to <i>Erythrosuchus </i>(as foreseen in Woodward's postscript). The large pedal ungual I with obliquely curved unguals was compared favorably to sauropods, but is also present in derived hyperodapedontines. Finally, a pedal digit with four phalanges was considered similar to dinosaurs and unlike 'anomodonts', but rhynchosaurs have three pedal digits with this many phalanges as well, and the digit closely matches manual digit III of Alemoa <i>Hyperodapedon</i>. Woodward's 1907 paper was actually written in 1904, and when reprinted in 1908 he included a postscript which recognized BMNH R2791 as non-dinosaurian. As he compared it favorably to <i>Erythrosuchus </i>(considered by Woodward to resemble both 'anomodonts' and 'belodonts'- the latter containing parasuchians and aetosaurs), Woodward now considered <i>Scaphonyx </i>an 'anomodont'. <br />Huene (1908) noted <i>Scaphonyx </i>was unlike dinosaurs in the presence of postaxial intercentra, cervical diapophyses and parapophyses which are placed high on the vertebra, and dissimilar unguals. He suggested it might be a therapsid or parasuchian. In 1911, Huene proposed <i>Scaphonyx </i>and <i>Erythrosuchus </i>were members of his new 'thecodont' group <i>Pelycosimia</i>, which continued through 1926 when he gave <i>Scaphonyx </i>its own family. In 1929, Huene finally recognized the similarity between <i>Scaphonyx </i>and rhynchosaurs, assigning the genus to the group.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-ushn3vsq5Zs/Vfpvw9UeF3I/AAAAAAAAAsk/v4i8P-MUbpg/s1600/Scaphonyx%2Bfischeri.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="249" src="http://3.bp.blogspot.com/-ushn3vsq5Zs/Vfpvw9UeF3I/AAAAAAAAAsk/v4i8P-MUbpg/s320/Scaphonyx%2Bfischeri.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Scaphonyx fischeri</i> (BM R-5033).&nbsp; Upper left- posterior cervical centrum in anterior and right lateral view.&nbsp; Upper right- anterior dorsal centrum in anterior and right lateral view.&nbsp; Lower left- ?manual digit III in dorsal and ventral view, phalanx III-1 is rotated 90 degrees. Lower right- pedal ungual I in medial, proximal and lateral view.&nbsp; All to scale (after Woodward, 1908).</td></tr></tbody></table>While long considered a valid genus of rhynchosaur, Langer (1996; published in Langer and Schultz, 2000a) proposed <i>Scaphonyx fischeri</i>'s holotype is indeterminate, as multiple species are known from the Alemoa member (<i>mariensis</i>, <i>sanjuanensis</i>, and what would be named <i>huenei</i>) which have only been distinguished using cranial characters. Although <i>huenei </i>is not known from postcrania (so can't be compared to <i>fischeri</i>), <i>mariensis </i>and <i>sanjuanensis </i>have not had their vertebral or pedal anatomy compared in detail in the published literature. Indeed, Alemoa rhynchosaurs have not had their postcrania well described in over seventy years. Given these facts and that I lack access to both Langer's thesis and <i>mariensis</i>' original and only published description, I only consider it provisionally indeterminate here. Additional specimens assigned to <i>S. fischeri</i> by Huene (1926, 1942) have been considered indeterminate or referrable to <i>S. sanjuanensis</i> (Langer and Schultz, 2000b; Montefeltro, 2008; Langer, pers. comm. 2015).<br /><br /><b>References</b>- Woodward, 1903. On some dinosaurian bones from south Brazil. Geological Magazine. 10(11), 512.<br />White, 1906. Geology of south Brazil. Science. 24(612), 377-379.<br />Woodward, 1907. On some fossil reptilian bones from the state of Rio Grande do Sul. Revista do Museu Paulista. 7, 46-57.<br />Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419. <br />Woodward, 1908. On some fossil reptilian bones from the state of Rio Grande do Sul. Geological Magazine. 5(6), 251-255.<br />Huene, 1911. Über <i>Erythrosuchus</i>, Vertreter der neuen Reptil-Ordnung Pelycosimia. Geologische und Paläontologische Abhandlungen. 10(1), 1-60. <br />Huene, 1926. Gondwana-Reptilien in Südamerika. Palaeontologia Hungarica. 2, 1-108.<br />Huene, 1929. Über Rhynchosaurier und andere Reptilien aus den Gondwana-Ablagerungen Südamerikas. Geologie und Palaeontogie Abhandlungen. 17, 1-61.<br />Huene, 1942. Die fossilen Reptilien des sudamerikanischen Gondwanalandes. C. H. Beck, Munich. 342 pp.<br />Langer, 1996. Rincossauros sul-brasileiros: Historico e filogenia. Masters thesis, Universidade Federal do Rio Grande do Sul. 361 pp.<br />Langer and Schultz, 2000a. Rincossauros-herbivoros cosmopolitas do Triassico. In Holz and de Ros (eds.). Paleontologia do Rio Grande do Sul. Porto Alegre. Ediitora da Universidade, CIGO/UFRGS, Brazil. 246-272. <br />Langer and Schultz, 2000b. A new species of the Late Triassic rhynchosaur <i>Hyperodapedon </i>from the Santa Maria Formation of south Brazil. Palaeontology. 43, 633-652. <br />Whatley, 2005. Phylogenetic relationships of <i>Isalorhynchus genovefae</i>, the rhynchosaur (Reptilia, Archosauromorpha) from Madagascar. PhD thesis, University of California. 276 pp.<br />Montefeltro, 2008. Inter-relações filogenéticas dos rincossauros (Diapsida, Archosauromorpha). Masters thesis, Universidade de Sao Paulo. 203 pp.<br /><br /><br /><br />As a bonus, since I make a short entry for every species of a genus that I use on The Theropod Database, I came across <i>Hyperodapedon minor</i>.&nbsp; Barely any info was present online, or in the literature.<br /><br /><i><b>H. gordoni</b></i> Huxley vide Murchison, 1858<br />?= <i>Hyperodapedon minor</i> Burckhardt, 1900b<br />= <i>Stenometopon taylori</i> Boulenger, 1904<br /><b>Early Norian, Late Triassic<br />Lossiemouth Sandstone Formation, Scotland<br />Comments</b>- The citation for Murchison (1858) is often listed incorrectly, misspelled 'Murchinson', cited as 1859, with an erroneous title, and pagination from Huxley's 1869 paper.<br /><i><b>Hyperodapedon minor</b></i>- This species was established by Burckhardt (1900b; page 492) for two small maxillae and a mandible from Warwickshire which were mentioned in a footnote by Huxley (1869) as <i>H. gordoni</i>. Only a few statements were made about <i>H. minor</i> in Burckhardt's work, with the only proposed distinguishing character being a more posteriorly extensive dentary tooth row than <i>H. gordoni</i>. The taxon has been virtually ignored in the literature since, though Huene (1942) did say its distinctiveness from <i>H. gordoni </i>is unfounded and that it should probably be rejected. Discussion with Benton (pers. comm 2015) indicates Burckhardt only visited the BMNH, though no specimens there were indicated as belonging to this species. Based on Benton's unpublished thesis notes, I believe BMNH R3150 (listed as "Partial skull 18 pieces, some fitting: palate views of mx, pal etc - small animal") is the best possibility for being <i>H. minor</i>'s holotype, though it's possible the holotype has remained unnoticed or become lost. Regardless, the fact all diagnostic Lossiemouth Sandstone rhynchosaurs have been referred to <i>H. gordoni</i> suggests <i>H. minor</i> is similarly referrable.<br /><b>References</b>- Murchison, 1858. On the sandstones of Morayshire (Elgin, &amp;c.) containing reptilian remains; and on their relations to the Old Red Sandstone of that country. Quarterly Journal of the Geological Society of London. 15, 419-439.<br />Huxley, 1869. On <i>Hyperodapedon</i>. Quarterly Journal of the Geological Society of London. 25, 138-152.<br />Burckhardt, 1900a. On <i>Hyperodapedon gordoni</i>. Geological Magazine. 7(12), 529-535. <br />Burckhardt, 1900b. On <i>Hyperodapedon gordoni</i>. Geological Magazine. 7(37), 486-492.<br />Boulenger, 1904. On reptilian remains from the Triass of Elgin. Philosophical Transactions of the Royal Society of London B. 196, 175-189.<br />Huene, 1942. Die fossilen Reptilien des sudamerikanischen Gondwanalandes. C. H. Beck, Munich. 342 pp. <br />Benton, 1981. The Triassic reptile <i>Hyperodapedon </i>from Elgin, functional morphology and relationships. PhD thesis, University of Newcastle upon Tyne. [? pp]. Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com0tag:blogger.com,1999:blog-3248412803814730250.post-39243300836454945612015-09-01T00:17:00.001-07:002015-09-02T00:02:54.152-07:00Plesiosaurs that used to be dinosaursSeveral marine reptiles have been confused for dinosaurs in the past, from sea lizard <a href="http://theropoddatabase.com/Non-theropods.htm#Taniwhasaurusmikasaensis" target="_blank">"Yezosaurus"</a>, to ichthyosaur <a href="http://theropoddatabase.com/Non-theropods.htm#Rachitremapellati" target="_blank"><i>Rachitrema</i></a>, to sea turtle <a href="http://theropoddatabase.com/Non-theropods.htm#Pneumatoarthruspeloreus" target="_blank"><i>Pneumatoarthrus</i></a>.&nbsp; But what about plesiosaurs?&nbsp; These are the specimens that I know of which have been previously assigned to saurischians.&nbsp; If this has taught me anything, it's that there are a TON of century-old plesiosaur taxa that need to be reexamined.<br /><br /><b>unnamed thalassophonean</b> (Hahnel, 1988)<br /><b>Middle Kimmeridgian, Late Jurassic<br />La Caja Formation, Mexico<br />Material</b>- (UANL-FCT-R2; The Monster of Aramberri) (~15 m) jaw fragment (lost), cranial fragments, nine cervical vertebrae, seven partial pectoral vertebrae (90-105 mm), rib fragments, gastralium?, axial material, incomplete scapula, incomplete coracoids, humerus, pelvis, femora (~1.2 m), epipodials<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-AuuSPcqxAZw/VeTt_09xXxI/AAAAAAAAAqU/iynhR6womkI/s1600/LJ%2BMexico%2Bplesiosaur%2Bsnout%2Bnon%2Btheropod.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="213" src="http://1.bp.blogspot.com/-AuuSPcqxAZw/VeTt_09xXxI/AAAAAAAAAqU/iynhR6womkI/s320/LJ%2BMexico%2Bplesiosaur%2Bsnout%2Bnon%2Btheropod.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Jaw fragment of thalassophonean UANL-FCT-R2 which caused Hahnel to mistake it for a theropod (after Buchy, 2007).</td></tr></tbody></table><br /><b>Comments</b>- Hahnel (1988) initially referred this specimen to Theropoda based on the large size and carnivorous teeth. Buchy et al. (2003) reidentified it as Pliosauridae indet. based on the pectoral section and lost snout. Frey et al. (2006) announced the recovery of much of the rest of the specimen, preliminary results which are given in Buchy's (2007) thesis. Buchy retained it as Pliosauridae indet., which can be narrowed to Thalassophonea indet. given its late age. Once more of the specimen is prepared, it may be possible to assign further.<br /><b>References</b>- Hahnel, 1988. Hallazgo de restos de dinosaurio en Aramberri, N.L., Mexico. Actas de la. Facultad de Ciencias de la Tierra, U.A.N.L. 3, 245-250.<br />Buchy, Frey, Stinnesbeck and Lopez-Oliva, 2003. First occurrence of a gigantic pliosaurid plesiosaur in the Late Jurassic (Kimmeridgian) of Mexico. Bulletin de la Societe Geologique de France. 174(3), 271-278.<br />Frey, Stinnesbeck and Buchy, 2006. The Monster of Aramberri. German Research. 27(3), 4-7.<br />Buchy, 2007. Mesozoic marine reptiles from north-east Mexico: Description, systematics, assemblages and palaeobiogeography. PhD thesis, Universitat Karlsruhe. 89 pp. <br /><br /><b>unnamed probable brachauchenine</b> (Knoll, Collete, Dubus and Petit, 2000)<br /><b>Early Albian, Early Cretaceous<br />Sables verts, France<br />Material</b>- (Petit coll.) posterior dorsal centrum (107 mm)<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-GWLfyouZhzo/VeUeQU9GkFI/AAAAAAAAAqo/nZzpsQNEC9w/s1600/EC%2BFrance%2Bpliosaur%2Bvertebra%2Bnon-sauropod.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://1.bp.blogspot.com/-GWLfyouZhzo/VeUeQU9GkFI/AAAAAAAAAqo/nZzpsQNEC9w/s320/EC%2BFrance%2Bpliosaur%2Bvertebra%2Bnon-sauropod.jpg" width="299" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Posterior dorsal centrum (Petit coll.) of a ?brachauchenine pliosaurid first published as a sauropod caudal by Knoll et al., in anterior (A), left lateral (B), posterior (C) and ventral (D) views (after Buffetaut et al., 2005).</td></tr></tbody></table><br /><b>Comments</b>- Knoll et al. (2000) originally described this as a possibly brachiosaurid sauropod proximal caudal, but it was reidentified by Buffetaut et al. (2005) as a pliosaurid dorsal. Given its age, it is probably referrable to Brachaucheninae, but is likely to be indeterminate.<br /><b>References</b>- Knoll, Collete, Dubus and Petit, 2000. On the presence of a sauropod dinosaur (Saurischia) in the Albian of Aube (France). Geodiversitas. 22, 389-394.<br />Buffetaut, Collete, Dubus and Petit, 2005. The "sauropod" from the Albian of Mesnil-Saint-Père (Aube, France): A pliosaur, not a dinosaur. Carnets de Géologie. 2005/01, 1-5.&nbsp; <br /><br /><i><b>Pliosaurus </b></i>Owen, 1841 sensu Owen, 1842<br />?= <i>Spondylosaurus </i>Fischer de Waldheim, 1845<br />?= "Tapinosaurus" Lennier, 1887 <br />= <i>Stretosaurus </i>Tarlo, 1959<br />= <i>Strongylokrotaphus </i>Novozhilov, 1964<br /><b>Comments</b>- This was originally erected as a subgenus of <i>Plesiosaurus</i>, spelled <i>Pleiosaurus </i>(Owen, 1841). As Benson et al. (2013) state, ICZN Article 33.3.1 indicates <i>Pliosaurus </i>is an incorrect subsequent spelling but should be retained as it has been "in prevailing usage and is attributed to the publication of the original spelling." Owen (1841) raised it to genus level. See Knutsen (2012) and Benson et al. (2013) for differing views on species validity.<br /><b>References</b>- Owen, 1841. Odontography; or a treatise on the comparative anatomy of the teeth, I Part 11. Dental system of reptiles. Hippolyte Bailliere, London. 179-295.<br />Owen, 1842. Report on British fossil reptiles. Part II. Report of the Eleventh Meeting of the British Association for the Advancement of Science. 60-204.<br />Fischer de Waldheim, 1845. Notice sur le <i>Spondylosaurus </i>genre de saurien fossile de l'oolithe de Moscow. Aus dem Bulletin de la Societe Imperiale des Naturalistes de Moscou. 18, 343-351. <br />Lennier, 1887. Études paléontologiques. Description des fossiles du Cap de la Hève. Bulletin de la Société Géologique de Normandie. 1886(12), 17-98. <br />Tarlo, 1959. <i>Stretosaurus </i>gen. nov., a giant pliosaur from the Kimmeridge Clay. Palaeontology. 2, 39-55.<br />Knutsen, 2012. A taxonomic revision of the genus <i>Pliosaurus </i>(Owen, 1841a) Owen, 1841b. Norwegian Journal of Geology. 92, 259-276. <br />Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013. A giant pliosaurid skull from the Late Jurassic of England. PLoS ONE. 8(5), e65989.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-_GupnWrLIPM/VeUiGiY_wbI/AAAAAAAAAq4/VVVgLr0apYY/s1600/Spondylosaurus.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="118" src="http://1.bp.blogspot.com/-_GupnWrLIPM/VeUiGiY_wbI/AAAAAAAAAq4/VVVgLr0apYY/s320/Spondylosaurus.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Not ever thought to be a dinosaur, but how often do you get to see the holotype of <i>Spondylosaurus frearsi</i>?&nbsp; This probable <i>Pliosaurus </i>specimen from the Late Jurassic of Russia hasn't been redescribed in 170 years, and according to Storrs et al. (2000) is of unknown whereabouts.&nbsp; Posterior cervical centrum in (left to right) ventral, left lateral, anteroventral and oblique anterodorsolateral views (after Fischer de Waldheim, 1845).</td></tr></tbody></table><br /><i><b>P. rigauxi</b></i> (Sauvage, 1874) new comb.<br />= <i>Cetiosaurus rigauxi</i> Sauvage, 1874<br /><b>Middle Tithonian, Late Jurassic<br />Portel, France<br />Holotype</b>- (MHNL coll.; = MHNB 233) posterior cervical centrum (85 mm)<br /><br /><b>Comments</b>- Sauvage (1874) initially described this as a posterior cervical centrum of a new <i>Cetiosaurus </i>species, notable for its short proportions. In 1895 he referred it to <i>Pliosaurus sp</i>. in a brief note, which would technically make it <i>Pliosaurus rigauxi</i>. That combination has never been used to my knowledge, however. Sauvage (1902) later referred the specimen to <i>Pliosaurus grandis</i> without stated justification. <i>P? grandis</i> is based on a scapula and three unassociated propodials from the Kimmeridgian of England which were poorly described and lost, so there's no reason to connect them to <i>rigauxi</i>. No more recent studies have been published, and the specimen has never been illustrated. The specimen number is taken from Fossilworks' website, though the Museum d'Histoire naturelle de Boulogne-sur-Mer (MHNB) closed in 2003 and transferred its collections to the Musée d'Histoire naturelle de Lille. Thus <i>rigauxi </i>presumedly has a MHNL number now instead.<br /><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-sc8Qv1ziMOg/VeUtTUQkvlI/AAAAAAAAArM/cRCXgu5hxUA/s1600/Pliosaurus%2Bsuprajurensis.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="154" src="http://4.bp.blogspot.com/-sc8Qv1ziMOg/VeUtTUQkvlI/AAAAAAAAArM/cRCXgu5hxUA/s320/Pliosaurus%2Bsuprajurensis.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Pliosaurus suprajurensis</i>, incomplete <strike>anterior cervical</strike> pectoral (thanks Sven Sachs) vertebra (MHNL coll.) in (left to right) ventral, left lateral, and anterior views (after Sauvage, 1879).&nbsp; This may be the same species as <i>"Cetiosaurus" rigauxi</i>, though the latter has never been figured.</td></tr></tbody></table><br />Based on the original description, <i>rigauxi </i>differs from sauropods such as <i>Cetiosaurus </i>in the short centrum (52% of height, compared to no less than 154% in <i>Cetiosaurus</i>), with even the short-necked <i>Brachytrachelopan </i>having subequal proportions at best. Furthermore, all gravisaurs have strongly opisthocoelous cervicals, whereas <i>rigauxi</i>'s is weakly amphicoelous. The parapophyses are 55% of centrum height, while they are much smaller in sauropods (e.g. 34% in cervical 12 of <i>Cetiosaurus</i>). All of these features are seen in pliosaur posterior cervicals however (e.g. length/width ratio of 51-58% and parapophysis/centrum height ratio 70% in <i>P. brachydeirus</i>' holotype). The undivided parapophysis located on the centrum indicates Sauvage had its position in the vertebral column correct despite assigning it to the wrong group. Notably, Sauvage states the ventral surface is "cut into a peak", which suggests a median keel like that which characterizes <i>P. brachydeirus</i>. The latter species' holotype is from the Early Kimmeridgian, but at least one other keeled pliosaur centrum is known from the Tithonian of France- <i>Pliosaurus suprajurensis</i>. It's possible both <i>rigauxi </i>and <i>suprajurensis </i>are members of a long-lived <i>P. brachydeirus</i>, or that the French species are synonymous but distinct from <i>P. brachydeirus</i>. In the latter case, <i>rigauxi </i>would have priority over <i>suprajurensis</i>, but they cannot be directly compared as they're from different sections of the neck (though <i>rigauxi </i>is from a much larger individual, with length x height x width 85x165x185 mm vs. 50x68x75 mm in <i>suprajurensis</i>), and its likely neither can be anatomically distinguished from <i>P. brachydeirus</i>. <i>rigauxi </i>is not officially synonymized here though, pending verification of the keeled morphology and modern description of the specimen.<br /><br /><b>References</b>- Sauvage, 1874. Mémoire sur les dinosauriens et les crocodiliens des terrains jurassiques de Boulogne-sur-Mer. Mémoires de la Société Géologique de France, série 2. 10(2), 1-57.<br />Sauvage, 1895. Les dinosauriens du terrain jurassique supérieur du Boulonnais. Bulletin de la Société Géologique de France, 3e série. 22, 465-470.<br />Sauvage, 1902. Note sur quelques Reptiles du Jurassique supérieur du Boulonnais. Bulletin de la Societe Academique de l‘Arrondissement de Boulogne-sur-Mer. 6, 380-398.<br />Fossilworks <a href="http://fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&amp;taxon_no=65160" target="_blank">http://fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&amp;taxon_no=65160&nbsp;</a><br /><br /><b><i>P? sp.</i></b> (Lennier, 1887)<br />= "Tapinosaurus" Lennier, 1887<br /><b>Late Kimmeridgian, Late Jurassic<br />lower Argiles d'Ecqueville Member of the Argiles d'Octeville Formation, France<br />Material</b>- (Muséum d’Histoire naturelle du Havre; destroyed) anterior cervical centrum (70 mm) (Lennier, 1887) <br />(Muséum d’Histoire naturelle du Havre; destroyed) three cervical centra (80, 78, 65 mm), two cervical ribs, two partial cervical or anterior dorsal neural arches, four incomplete dorsal neural arches, six dorsal ribs (four partial, one fragmentary; 1 m) (Rabeck, 1925)<br />?...(Lepage coll. 15.8.31.E1) posterior cervical centrum (91 mm) (Lepage et al., 2009)<br />?...(Muséum du Havre IIFD358) incomplete dorsal rib (Lepage et al., 2009)<br /><b>Early Kimmeridgian, Late Jurassic<br />Marnes de Bleville, France</b><br />(Muséum d’Histoire naturelle du Havre; destroyed) proximal dorsal rib (Lennier, 1887) <br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-yg7PCeCVdlE/VeU9XktuzOI/AAAAAAAAArg/_2vEZW2MeJE/s1600/Tapinosaurus.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" src="http://1.bp.blogspot.com/-yg7PCeCVdlE/VeU9XktuzOI/AAAAAAAAArg/_2vEZW2MeJE/s1600/Tapinosaurus.jpg" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>?Pliosaurus </i>anterior cervical centrum (Muséum d’Histoire naturelle du Havre coll.) in anterior or posterior view first figured as <i>Tapinosaurus sp.</i>, a misspelling of <i>Tapinocephalus </i>(after Lennier, 1887).</td></tr></tbody></table><br /><b>Comments</b>- Lennier (1887) referred an unassociated cervical centrum and dorsal rib (both originally in the Muséum d’Histoire naturelle du Havre, but destroyed in 1944) to <i>Tapinocephalus</i>, then thought to be dinosaurian but now known to be a dinocephalian synapsid. This was seemingly due to their large size. The figure caption of Lennier's plate illustrating the centrum mistakenly said "<i>Tapinosaurus sp</i>?", clearly a misspelling of <i>Tapinocephalus </i>and not previously suggested to be a valid genus. <br />Rabeck (1925) described a specimen found in 1923 as the dinosaur "<i>Tapinosaurus sp</i>?", based on resemblence to Lennier's specimens, unaware that "Tapinosaurus" was not an accepted genus. This specimen (consisting of partial vertebrae and ribs) was also held at the Muséum d’Histoire naturelle du Havre and similarly destroyed in WWII. Stiegelmann (1925) provided measurements of the specimen.<br /><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-zgZDcdU-A_g/VeVIAwQZhUI/AAAAAAAAAr0/GskNHUmLaCU/s1600/Tapinosaurus%2BRabeck.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="118" src="http://3.bp.blogspot.com/-zgZDcdU-A_g/VeVIAwQZhUI/AAAAAAAAAr0/GskNHUmLaCU/s320/Tapinosaurus%2BRabeck.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Only existing photo of Rabeck's "<i>Tapinosaurus sp.</i>" material (Muséum d’Histoire naturelle du Havre coll.), consisting of three cervical centra (left center), two cervical ribs (around the bottom centrum), neural arches and dorsal ribs (after Lepage et al., 2009).</td></tr></tbody></table><br />After this, "Tapinosaurus" was seldomly mentioned. Kuhn (1939) includes Rabeck's material questionably under <i>Omosaurus</i>, which is treated as Saurischia indet. by Kuhn despite being stegosaurid (a replacement name for <i>Dacentrurus</i>). Steel (1970) realized Lennier's article was supposed to reference <i>Tapinocephalus</i>, but stated Rabeck's material "seemingly pertains to a large dinosaur, but is indeterminable", placing "Tapinosaurus" in Sauropoda incertae sedis.&nbsp; Rabeck's specimen is not a sauropod, as it has short amphicoelous cervical centra without pleurocoels, short laterally oriented cervical ribs, dorsal neural arches lacking laminae, and single-headed dorsal ribs.&nbsp; Buffetaut et al. (1991) first noticed the discrepency between the article and plate caption in Lennier's work, and identified both this centrum and Rebeck's "Tapinosaurus" as sauropterygians. Similarly, Molnar (pers comm. in Olshevsky, 1991) stated these specimens were probably plesiosaurian. Finally, Lepage et al. (2009) published detailed overview of "Tapinosaurus"' history (forming the basis of most of this entry), and redescribed the specimens as <i>Pliosaurus sp.</i> for Lennier's and <i>Pliosaurus </i>cf. <i>macromerus</i> for Rabeck's. They also described three new specimens from the lower Argiles d'Ecqueville, two of which might be referrable to the same individual as Rabeck's specimen as they are from the same layer and of similar size.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-ydCrFp5lrsA/VeVM_oBgkjI/AAAAAAAAAsI/wfhVgy8GY5c/s1600/Pliosaurus%2Barchiaci.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="178" src="http://3.bp.blogspot.com/-ydCrFp5lrsA/VeVM_oBgkjI/AAAAAAAAAsI/wfhVgy8GY5c/s320/Pliosaurus%2Barchiaci.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Pliosaurus archiaci</i> (MNHN 24.1), dentaries in dorsal and lateral views (after Lennier, 1870).&nbsp; This may be conspecific with the "Tapinosaurus" specimens.</td></tr></tbody></table><br /><b>"Tapinosaurus" is <i>Pliosaurus</i>?</b>- Unfortunately, the alpha level taxonomy of <i>Pliosaurus </i>is currently controversial, and most proposed distinguishing characters are cranial. The only axial character currently used to distinguish <i>Pliosaurus </i>species is the median ventral keel on cervical centra of <i>P. brachydeirus</i>. Rabeck's specimen lacks this, as do <i>P. brachyspondylus</i> (both current and proposed neotypes), <i>P. funkei</i> (paratype), <i>P. macromerus</i> (lectotype), <i>P. rossicus</i> (holotype) and <i>P. westburyensis</i>. Yet this morphology is plesiomorphic, also being found in e.g. <i>Brachauchenius</i>, <i>Simolestes </i>and <i>"P." andrewsi</i>. <i>Pliosaurus archiaci</i> is based on a mandible (MNHN 24.1) also discovered in the Kimmeridgian deposits of Le Havre, but cannot be compared to "Tapinosaurus". Another method would be to correlate the "Tapinosaurus" specimens stratigraphically with known <i>Pliosaurus </i>species. According to Lepage et al., at least Rabeck's specimen derives from the <i>Aulacostephanus mutabilis</i> zone of the Late Kimmeridgian. This corresponds to CAMSM J35990, a partial skeleton initially referred to a broad concept of <i>P. macromerus</i> (<i>Pliosaurus </i>without ventral cervical keels) but more recently found to be closest to <i>P. kevani</i> (in an analysis that did not include the <i>P. macromerus</i> lectotype or proposed neotype). CAMSM J35990 is similar Rabeck's specimen in being larger than most and lacking ventral cervical keels, so there may be a real large <i>A. mutabilis</i> zone species of <i>Pliosaurus </i>that is currently undiagnosed. This may correspond to the large <i>P. portentificus</i>, although that has been considered a nomen dubium and may belong to the more recent <i>A. euxodus</i> zone. Perhaps notable is that <i>P. portentificus</i> has the same number of symphyseal alveoli (8) as <i>P. archiaci</i>, whereas other species have more (&gt;11 in <i>P. brachydeirus</i>, 9 in <i>P. carpenteri</i>, ~14-15 in <i>P. kevani</i>) or less (6 in <i>P. rossicus</i> and <i>P. patagonicus</i>). In any case, both <i>P. macromerus</i>' lectotype and proposed neotype are from more recently deposited sediments, so Lepage et al.'s assignment seems unlikely. Pending further studies, the "Tapinosaurus" material is best retained as <i>Pliosaurus sp.</i>. Lennier's dorsal rib is from a different locality, whose age corresponds to <i>P. brachydeirus</i> and <i>P. kevani</i>, though it is near certainly indeterminate.<br /><br /><b>References</b>- Lennier, 1887. Études paléontologiques. Description des fossiles du Cap de la Hève. Bulletin de la Société Géologique de Normandie. 1886(12), 17-98.<br />Rabeck, 1925. Notes sur la découverte d'ossements de dinosaurien dans les Argiles supérieures Kimméridgiennes du Cap de la Hève (Octeville-sur-Mer). Bulletin de la Société Géologique de Normandie. 1916/1923(34), 72-74<br />Stiegelmann, 1925. Note additionnelle [à Notes sur la découverte d'ossements de dinosaurien dans les Argiles supérieures Kimméridgiennes du Cap de la Hève (Octeville-sur-Mer) de G. Rabeck]. Bulletin de la Société Géologique de Normandie.1916/1923(34), 75.<br />Kuhn, 1939. Saurischia. In Fossilium Catalogus I. Animalia. 87. 124 pp.<br />Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp. <br />Buffetaut, Cuny and Le Loeuff, 1991. French dinosaurs: The best record in Europe? Modern Geology. 16, 17-42.<br />Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp. <br />Lepage, Buffetaut and Lepage, 2009. Qu'est-ce que <i>Tapinosaurus</i>? Lennier, Rabeck et les grands Sauroptérygiens du Kimméridgien supérieur de la région havraise (Normandie, France). Bulletin de la Société géologique de Normandie et des amis du Muséum du Havre. 96(1), 27-59.<br /><br /><b>Blog entry specific references</b>- Lennier, 1870. Etudes géologiques et paléontologiques sur l'embouchure de la Seine et les Falaises de la Haute-Normandie. Imprimerie Eugène Costey, Havre. 245 pp.<br /><br />Sauvage, 1879. Prodrome des Plesiosauriens et des Elasmosauriens des formations Jurassiques superieures de Boulogne-sur-Mer. Annales des Sciences Naturelles, 6 Serie. 8(13), 1-38.<br /><br />Storrs, Arkhangelskii and Efimov, 2000. Mesozoic marine reptiles of Russia and other former Soviet Republics. In Benton, Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press. 187-210.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com2tag:blogger.com,1999:blog-3248412803814730250.post-76745641562690284582015-08-25T17:43:00.001-07:002015-08-25T17:43:44.875-07:00Another Triassic tooth, Paleosaurus fraserianusToday is the last day of my three week vacation, during which I got a lot accomplished.&nbsp; This included adding four ex-saurischians to the Theropod Database, which will each get their own post this week before I upload this month's updates to the website.&nbsp; The first of these ex-saurischians is one whose identity was solved a century ago, but continues to be misunderstood by dinosaur workers.<br /><br /><i><b>Clepsysaurus? fraserianus</b></i> (Cope, 1878a) Hay, 1930<br />= <i>Paleosaurus fraserianus</i> Cope, 1878a (as <i>Palaeosaurus fraserianus</i>)<br />= <i>Thecodontosaurus fraserianus</i> (Cope, 1878a) Hay, 1902<br />= <i>Palaeosauriscus fraserianus</i> (Cope, 1878a) Kuhn, 1965<br /><b>Norian, Late Triassic<br />New Oxford Formation, Pennsylvania, US<br />Holotype</b>- (AMNH 1861) tooth (20x6.5x? mm)<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-Ct0T0NXY-aY/Vdz8A8y1EFI/AAAAAAAAAp4/vm9FyfpJbd0/s1600/fraserianus%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="108" src="http://4.bp.blogspot.com/-Ct0T0NXY-aY/Vdz8A8y1EFI/AAAAAAAAAp4/vm9FyfpJbd0/s320/fraserianus%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Paleosaurus fraserianus</i> (AMNH 1861) at center with section (after Huene, 1921) compared to from left to right- holotype of <i>Paleosaurus cylindrodon</i> (BRSMG Ca7449/4) with section (after Huxley, 1870 and Huene, 1908 respectively); right fourth premaxillary tooth of <i>Nicrosaurus kapffi</i> (SMNS 13078) with sections (after Hungerbuhler, 2000); eighth and ninth dentary teeth of <i>Thecodontosaurus antiquus</i> neotype (BRSMG Ca4529/2) (after Galton, 2007); third dentary tooth of <i>Anchisaurus polyzelus</i> (YPM 209) with drawing (after Fedak and Galton, 2007).</td></tr></tbody></table><br /><br /><b>Comments</b>- Cope (1878a) described this tooth (which was first presented the year prior) as a new species of <i>Palaeosaurus</i>, at the time a common misspelling of <i>Paleosaurus</i>. Olshevsky (2000) was the first to correct the genus' spelling in this binomial. Note there is a valid genus <i>Palaeosaurus </i>(Geoffroy Sant-Hillaire, 1836; which is currently a junior synonym of the teleosaurid <i>Steneosaurus</i>) however, which caused Kuhn (1965) to incorrectly think <i>Paleosaurus </i>was preoccupied since its spelling is so similar. Thus he referred all <i>Paleosaurus </i>species to his new genus <i>Palaeosauriscus</i>, but this is unnecessary according to the ICZN. <i>Paleosaurus </i>itself is based on <i>P. cylindrodon</i>, an archosauriform tooth of uncertain affinities from the Norian of England which differs from <i>fraserianus </i>in having elongate and oblique serrations, being less recurved, and having a more tapered distal edge in section (far left in figure above).<br /><br /><b><i>fraserianus </i>a dinosaur?</b> Nopsca (1901) was the first to assign the species explicitly to Dinosauria or Theropoda, assigning it to a subfamily Anchisauridae [sic] within Megalosauridae, but his anchisaurids consisted largely of basal sauropodomorphs and Triassic carnivorous archosauriform teeth. Hay (1902) had a similar concept for Anchisauridae within his Theropoda, similarly placing <i>fraserianus </i>there though assigning it to <i>Thecodontosaurus</i>, as he synonymized the genus with <i>Paleosaurus</i>. Note Colbert and Chaffee (1941) wrongly cited Cope (1878b) as using the combination <i>Thecodontosaurus fraserianus</i>, but that work only uses <i>Thecodontosaurus </i>for <i>T. gibbidens</i>. Hay (1930) retained <i>fraserianus </i>in Anchisauridae and Theropoda, but now placed it in the genus <i>Clepsysaurus </i>(a parasuchian), perhaps based on the similarity noted by Huene (see below). Steel (1970) referred it to his theropodan Ornithosuchidae, which besides <i>Ornithosuchus </i>contained <i>Teratosaurus</i>, Triassic carnivorous archosauriform teeth, and basal sauropodomorph remains incorrectly associated with the latter. The most confusing generic assignment has been that of Olshevsky (1991, 2000), who made <i>fraserianus </i>a junior synonym of <i>Anchisaurus polyzelu</i>s, which is from the much later (Pliensbachian) Portland Formation of Connecticut. fraserianus is quite unlike sauropodomorph teeth (including <i>Thecodontosaurus </i>and <i>Anchisaurus</i>; pictured at right in figure above) in being highly recurved, with an unconstricted base, little labiolingual compression, and small serrations which are perpendicular to the tooth axis. The connection was maintained through history largely via ignorance of <i>fraserianus</i>' actual morphology in addition to continued confusion of <i>Paleosaurus </i>with <i>Thecodontosaurus </i>and <i>Efraasia</i>.<br /><br /><b><i>fraserianus </i>a parasuchian?</b> Lesley (1889) may be the first author to suggest <i>fraserianus </i>is parasuchian, albeit without evidence. Huene (1921) has been the only author to illustrate <i>fraserianus</i>, and briefly described the specimen as well. Huene convincingly illustrated the similarity with other New Oxford parasuchian teeth. He suggested <i>fraserianus </i>was synonymous with <i>Clepsysaurus? veatleianus</i> and/or <i>Rutiodon carolinensis</i> from the same formation. Colbert and Chaffee (1941) made <i>fraserianus </i>a junior synonym of <i>Clepsysaurus pennsylvanicus</i>, based on geography. Of Norian archosauriforms which have recurved teeth with small serrations, only some proterochampsids and phytosaurid parasuchians are reported to have reduced labiolingual compression as in <i>fraserianus </i>(80% of FABL). While proterochampsid teeth remain largely undescribed, they are exclusively South American, so are an unlikely identification for <i>fraserianus</i>. Indeed, phytosaurid material is common in the New Oxford Formation, with <i>Rutiodon carolinensis</i> the only currently recognized valid taxon. It's therefore possible Huene was correct and that <i>fraserianus </i>is synonymous with <i>Rutiodon</i>, but the most recent review also suggested the presence of a larger, poorly characterized form (e.g. SMP VP-36; YPM-PU 11544). The tooth of <i>fraserianus </i>is much smaller than these latter elements, but could be ontogenetically young as well. Unfortunately, heterodonty is so great among phytosaurid dentitions, few of which have been described in detail, that it is not currently possible to assign isolated teeth to particular genera or species. Thus <i>fraserianus </i>remains Phytosauridae indet., and is here placed questionably in <i>Clepsysaurus</i> as that is the only phytosaurid genus prior authors have referred the species to.&nbsp; Based on the variation in <i>Nicrosaurus</i>, <i>fraserianus </i>may be based on a premaxillary or anterior maxillary tooth.<br /><br /><b>References</b>- Huxley, 1870. On the classification of the Dinosauria, with observations on the Dinosauria of the Trias. Quarterly Journal of the Geological Society of London. 26, 32-51.<br /><br />Cope, 1878a. On some saurians found in the Triassic of Pennsylvania, by C. M. Wheatley. Proceedings of the American Philosophical Society. 17(100), 231-232.<br /><br />Cope, 1878b. Triassic saurians from Pennsylvania. The American Naturalist. 12, 58. <br /><br />Lesley, 1889. A Dictionary of the Fossils of Pennsylvania and Neighboring States Named in the Reports and Catalogues of the Survey. Volume 2. The Board of Commissioners for the Geological Survey, Harrisburg. 914 pp.<br /><br />Nopcsa, 1901. A dinosaurusok atnezete es szarmazasa. Földtani Közlöny. 31, 193-224. <br /><br />Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America. United States Geological Survey Bulletin. 179, 868 pp.<br /><br />Huene, 1908. Die Dinosaurier der europäischen Triasformation mit Berücksichtiging der aussereuropäischen Vorkommnisse. Geologische und Paläontologische Abhandlungen Supplement-Band. 1, 419 pp. <br /><br />Huene, 1921. Reptilian and stegocephalian remains from the Triassic of Pennsylvania in the Cope collection. Bulletin American Museum of Natural History. 44(19), 561-574.<br /><br />Hay, 1930. Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution of Washington. 390(II), 1-1074.<br /><br />Colbert and Chaffee, 1941. The type of <i>Clepsysaurus pennsylvanicus</i> and its bearing upon the genus <i>Rutiodon</i>. <br /><br />Kuhn, 1965. Saurischia (Supplementum 1). In Fossilium Catalogus 1. Animalia. 109, 94 pp. <br /><br />Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.<br /><br />Hungerbuhler, 2000. <span class="st">Heterodonty in the European phytosaur <i>Nicrosaurus kapffi</i> and its implications for the taxonomic utility and functional morphology of phytosaur dentitions. Journal of Vertebrate Paleontology. 20(1), 31-48.</span><br /><br />Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings. 3, 1-157.<br /><br /> Fedak and Galton, 2007. New information on the braincase and skull of <i>Anchisaurus polyzelus</i> (Lower Jurassic, Connecticut, USA; Saurischia: Sauropodomorpha): Implications for sauropodomorph systematics. In Barrett and Batten (eds.). Evolution and Palaeobiology of Early Sauropodomorph Dinosaurs. Special Papers in Palaeontology. 77, 245-260.<span class="small"></span> <br /><br />Galton, 2007. Notes on the remains of archosaurian reptiles, mostly basal sauropodomorph dinosaurs, from the 1834 fissure fill (Rhaetian, Upper Triassic) at Clifton in Bristol, southwest England. Revue de Paléobiologie. 26(2), 505-591.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com2tag:blogger.com,1999:blog-3248412803814730250.post-46964052328367553072015-06-22T02:06:00.000-07:002015-06-22T02:06:46.466-07:00Chilesaurus brings out the BANDit in meIn the last post, I detailed what Novas et al.'s analyses actually indicated regarding <i>Chilesaurus</i>' relationships.&nbsp; Far from all indicating a basal tetanurine status, there was good support for positions in Sauropodomorpha, Coelurosauria or sister to Avepoda.&nbsp; What if we dig a bit deeper? <br /><br /><b>What kind of megalosaur looks like <i>Chilesaurus</i>?! </b><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-cyl2LCaa_Gw/VYdi3k39umI/AAAAAAAAAnA/W-TD05lzrVM/s1600/Chile%2BMono%2Bskull%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="175" src="http://1.bp.blogspot.com/-cyl2LCaa_Gw/VYdi3k39umI/AAAAAAAAAnA/W-TD05lzrVM/s320/Chile%2BMono%2Bskull%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Chilesaurus </i>(holotype SNGM-1935; premaxilla in medial view) and <i>Monolophosaurus </i>(IVPP V84019) snouts scaled to same premaxillary alveolar length (modified after Novas et al., 2015 and Brusatte et al., 2010 respectively).</td></tr></tbody></table><br />If you look at <i>Chilesaurus</i>, there are basically no characters similar to monolophosaur-piatnitzkysaur grade tetanurines.&nbsp; And I mean nothing.&nbsp; This is from the person who came up with a list of characters connecting <a href="http://theropoddatabase.blogspot.com/2010/09/ceratosaurian-ornithomimosaurs-another.html">ornithomimosaurs to ceratosaurs</a> and <a href="http://theropoddatabase.blogspot.com/2010/08/is-theropod-ornithischian-group-that.html">ornithischians to theropods</a>, so that's really saying something.&nbsp; The best I can do here is to note the scapula, humerus, radius and ulna wouldn't be out of place for a basal tetanurine, oh and it's coded as having a femoral extensor groove.&nbsp; I must admit I feel rather Feduccian saying this, but the result of placing <i>Chilesaurus </i>in Carrano et al.'s analysis looks to me like a case of cladistic failure where it has to land somewhere but the conclusion is unrealistic, being least terrible as opposed to most parsimonious.&nbsp; It's rather like when Chatterjee included <i>Protoavis </i>in his bird analyses- it emerged in a certain place (above <i>Archaeopteryx </i>but below ornithothoracines), but much of the morphology just didn't make sense there, from the unfused clavicles and plesiomorphic manus to the very derived pelvis.&nbsp; Another example is when alvarezsaurids were placed in Avialae, with their short coracoids, elongate chevrons, long ischia, etc.. <br /><br /><i>Chilesaurus </i>has no interdental plates, leaf-like teeth with marked wear facets, no postaxial epipophyses, no hyposphene-hypantrum articulations, no gastralia, a short manual phalanx II-2, one phalanx on manual digit III, no obturator process or foramen on its ischium, opisthopuby, separate pubic apices, no fibular crest, a tiny astragalar ascending process, huge pedal digit I which may contact the tarsus, completely unreduced proximal metatarsal III, etc..&nbsp; You'll see some of these in some tetanurines, but never close to so many at the same time (e.g. parvicursorines have up to 8, therizinosaurids have 4, ornithothoracines have up to 9).&nbsp; A few aren't even known to reverse (e.g. ascending process, fibular crest) regardless of the animal's ecotype.&nbsp; Even when the characters are common in theropods, they're not found at the megalosaur grade- e.g. elongate presacral centra with low neural spines, no metacarpal IV, no supracetabular crest, antitrochanter, supratrochanteric crest, small pubic peduncle, cylindrical anterior trochanter, etc..&nbsp; So immediately I'm suspicious.<br /><br /><b><i>Chilesaurus </i>tested in Coelurosauria </b><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-sAtltuZnOEY/VYdlZldO4-I/AAAAAAAAAnQ/J8B1UgTsLsg/s1600/Chile%2BPiat%2Bvert%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="138" src="http://3.bp.blogspot.com/-sAtltuZnOEY/VYdlZldO4-I/AAAAAAAAAnQ/J8B1UgTsLsg/s320/Chile%2BPiat%2Bvert%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Chilesaurus </i>(holotype SNGM-1935) and <i>Piatnitzkysaurus </i>(holotype PVL 4073; flipped horizontally) fourth cervical vertebrae in right lateral view, scaled to same centrum length minus anterior ball if present (modified after Novas et al., 2015 and Bonaparte, 1986 respectively).</td></tr></tbody></table><br />Noting most of these characters are found in coelurosaurs, which were not strongly rejected in the only matrix to include a vaguely representative sample (Smith et al.'s needed only 3 more steps to place it there), I added <i>Chilesaurus </i>to the Lori analysis (704 characters).&nbsp; <i>Monolophosaurus</i> is the outgroup, with some carnosaurs and basal coelurosaurs, and a complete sampling of maniraptoriforms minus derived ornithuromorphs. If it's really a basal tetanurine, <i>Chilesaurus </i>should end up down by <i>Monolophosaurus </i>due to the lack of characters present in various coelurosaurian groups.<br /><br />Without getting into the details of the topology, <i>Chilesaurus </i>emerged as a non-alvarezsaurid alvarezsauroid in a tree where <i>Haplocheirus </i>was closer to compsognathids as in several recent analyses.&nbsp; This makes a lot of sense compared to being a basal tetanurine, as it has the following characters shared with alvarezsaurids or more inclusive clades- *basally constricted teeth; *reduced tooth curvature; *no interdental plates; *no hyposphene-hypantra; *transversely compressed axial neural spine; *low presacral neural spines; *lower postaxial epipophyses; *amphicoelous cervical centra (as in <i>Alvarezsaurus</i>); *metacarpal I more than 66% mcII length; *manual phalanx II-2 shorter than II-1; reduced manual digit III; metacarpal IV absent; *low manual flexor tubercles; *poorly curved manual ungual I; pubic peduncle dorsoventrally shorter than ischial peduncle; *pubic peduncle short anteroposteriorly compared to ischial peduncle; *reduced supracetabular crest; *antitrochanter; *at least mesopuby; *no obturator foramen in pubis; *pubic apron reduced in length; *very small pubic expansion; *distal pubic expansions not combined medially (as in <i>Patagonykus</i>); *no ischial obturator process or foramen; *greater trochanter anteroposteriorly broader than femoral head; *cylindrical anterior trochanter; *proximally extensive anterior trochanter; *no anteromedial femoral crest distally; *cnemial crest with low anterior angle (as in patagonykines).<br /><br />Constraining <i>Chilesaurus </i>to be by <i>Monolophosaurus </i>is 13 steps longer, which is fairly well rejected.&nbsp; And the biostratigraphy fits, as alvarezsauroids diverged in the Middle Jurassic or earlier and (as <i>Haplocheirus </i>isn't one here) emerged in South America.&nbsp; Thus IF <i>Chilesaurus </i>is an avepod, I'd say a basal alvarezsauroid position is most likely.<br /><br /><b><i>Chilesaurus </i>tested in Ornithischia </b><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-cZrdP1syiwQ/VYdnfQ2C-rI/AAAAAAAAAnc/1fwMAHNhVZ8/s1600/Chile%2BMono%2Bpelvis%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="196" src="http://4.bp.blogspot.com/-cZrdP1syiwQ/VYdnfQ2C-rI/AAAAAAAAAnc/1fwMAHNhVZ8/s320/Chile%2BMono%2Bpelvis%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Chilesaurus </i>(paratype SNGM-1936) and <i>Monolophosaurus </i>(IVPP V84019; flipped horizontally) pelvis in right lateral view, scaled to same ilial length (modified after Novas et al., 2015 and Zhao and Currie, 1994 respectively).</td></tr></tbody></table><br />My second test is one Novas et al. should have done- code <i>Chilesaurus </i>in an ornithischian analysis.&nbsp; Most of the characters listed above as shared with alvarezsaurids are actually common in basal ornithischians, as indicated by asterisks.&nbsp; This brings to mind the hypothesis of Alifanov and Barsbold (2009) and Alifanov and Saveliev (2011) that alvarezsaurids aren't theropods and are more like ornithischians.&nbsp; As noted in my theropod-ornithischian post, the latter clade also has members with some of the supposed avepod characters of <i>Chilesaurus</i>- strap-like scapula, elongate preacetabular process, distal tibia backs fibula, etc..&nbsp; The jaws, manus, pelvis and pes of <i>Chilesaurus </i>all certainly look more ornithischian to me at a general level.&nbsp; I chose the newest version of Butler's ornithischian analysis, Han et al.'s (2012; 227 characters) with <i>Kulindadromeus </i>added as in Godefroit et al. (2014).&nbsp; In addition to adding <i>Chilesaurus</i>, I added <i>Monolophosaurus </i>to test Novas et al.'s hypothesis, along with <i>Allosaurus </i>as a complete non-coelurosaur tetanurine and standard theropod (the original matrix's only potential theropod is <i>Herrerasaurus</i>), Parvicursorinae and <i>Patagonykus </i>to test my above hypothesis, and <i>Haplocheirus </i>as an example of the kind of theropod alvarezsaurids evolved from.&nbsp; This also coincidentally tests Alifanov's idea.<br /><br />When run, <i>Chilesaurus</i> emerges as an iguanodont more derived than <i>Anabisetia</i>, but less than rhabdodontids, <i>Tenontosaurus </i>and dryomorphs.&nbsp; Alvarezsaurids emerge as theropods.&nbsp; But it only takes two more steps to make <i>Chilesaurus </i>sister to other ornithischians or a basal marginocephalian instead, and only three more to make it a basal cerapodan.&nbsp; Forcing it to be theropod takes 4 more steps, so still isn't too bad, and it comes out by alvarezsaurids.&nbsp; Forcing it to be by <i>Monolophosaurus </i>takes ten more steps, so seems unlikely.&nbsp; This might seem to indicate that <i>Chilesaurus </i>is plausibly an ornithischian, though without an obvious place in the clade (except probably not thyreophoran, where it takes 8 more steps).&nbsp; The taxa it usually ends up closest to are <i>Yueosaurus </i>and <i>Pisanosaurus</i>, the latter which is often found to be the most basal ornithischian.&nbsp; However, alvarezsaurids take only two steps to place sister to other ornithischians, and only five steps to place within Ornithischia (as thyreophorans or marginocephalians).&nbsp; Assuming alvarezsaurids are actually theropods, this suggests that the results of placing a theropod in an ornithischian analysis don't mean much.&nbsp; So while this test didn't reject an ornithischian <i>Chilesaurus</i>, I don't think it rejected an alvarezsauroid relationship strongly either.&nbsp;<br /><br />We also have the same problem as we did placing <i>Chilesaurus </i>in Alvarezsauroidea- it has features that are out of place for a non-basalmost Ornithischia and never or rarely seem to reverse.&nbsp; Premaxillary teeth extend to tip of element, no predentary, no cingulum (also in derived heterodontosaurines and derived iguanodonts), pleurocoels, no fourth manual digit, deep preacetabular process (also in some eurypodans), no prepubic process (also in <i>Pisanosaurus</i>), non-pendent fourth trochanter (also in pachycephalosaurs), unreduced fibular facet on astragalus, no ossified tendons along vertebrae (also in <i>Kulindadromeus</i>, <i>Koreanosaurus </i>and <i>Yueosaurus</i>).&nbsp; If <i>Pisanosaurus </i>is the basalmost ornithischian, it is more derived than <i>Chilesaurus </i>in having a reduced fibular facet on its astragalus yet less derived in lacking opisthopuby.&nbsp; Notably, Butler's matrix lacks any silesaurs, which may be relevant to basal Ornithischia if Langer and Ferigolo are right.<br /><br /><b><i>Chilesaurus </i>REtested in Sauropodomorpha</b><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-mHRiQF09qHE/VYeuzWRfkUI/AAAAAAAAAn0/-ggbWspQpLs/s1600/Chile%2BDilo%2Bpes%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="http://1.bp.blogspot.com/-mHRiQF09qHE/VYeuzWRfkUI/AAAAAAAAAn0/-ggbWspQpLs/s320/Chile%2BDilo%2Bpes%2Bcomp.jpg" width="231" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Chilesaurus </i>(paratype SNGM-1937; missing proximal ends of metatarsals I and II, distal digit IV, and metatarsal V) and <i>Dilophosaurus wetherilli </i>(holotype UCMP 37302; missing digit I and distal digit IV) pedes in proximal (top) and anterior (bottom) view, scaled to the same metatarsal III length (after Novas et al., 2015 and Welles, 1984 respectively).</td></tr></tbody></table><br />I next looked over <i>Chilesaurus</i>' codings in the Otero and Pol analysis that's based on Yates' sauropodomorph matrix.&nbsp; I found an appalling amount of un-coded and miscoded characters (45 of 361, or 12%).&nbsp; I list them here so that readers can see just how obvious many are, and that even if my interpretations of some are wrong, there are WAY too many to be excused.<br /><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>has a 'lateral plate' on its dentary (character 2), which is just to say the labial edge is much higher than the lingual edge, though it was coded as lacking one.</span><br /><span style="background-color: #f4cccc;">- It was left uncoded for its large subnarial premaxillary process (character 7).</span><br /><span style="background-color: #f4cccc;">- And its lack of a posteromedial premaxillary process (character 9).</span><br /><span style="background-color: #f4cccc;">- And the anterior narial edge being posterior to the center of the premaxillary ventral edge (character 18).</span><br /><span style="background-color: #f4cccc;">- And having the posterior narial border at least posterior to the premaxilla-maxilla suture (character 19).</span><br /><span style="background-color: #f4cccc;">- And lacking a strong inflection on the maxilla to form an obvious anterior ramus (character 25).</span><br /><span style="background-color: #f4cccc;">- And having a first dentary tooth adjacent to the symphysis (character 100).</span><br /><span style="background-color: #f4cccc;">- And having less than five premaxillary teeth (character 107).</span><br /><span style="background-color: #f4cccc;">- And having procumbant maxillary teeth (character 110).</span><br /><span style="background-color: #f4cccc;">- It was miscoded as lacking longitudinal labial grooves on its teeth (character 119), visible in the dentary teeth.</span><br /><span style="background-color: #f4cccc;">- Since Avepoda was coded as state 2 for character 147 ("Lateral surfaces of the dorsal centra: with invasive, sharp-rimmed pleurocoels") despite basally only having dorsal pleurocoels in anterior centra, and <i>Chilesaurus </i>is stated to have anterior dorsal pleurocoels, it is recoded to be state 2 as well.</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was oddly miscoded as lacking septate cervical pleurocoels (character 148), despite the emphasis on them in the figure and text.</span><br /><span style="background-color: #f4cccc;">- If the Nesbitt et al. matrix is correct, <i>Chilesaurus </i>is miscoded here as having hyposphene-hypantrum articulations (character 157).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>has transversely compressed dorsal neural spine cross sections (state 0 for character 169). </span><br /><span style="background-color: #f4cccc;">- The in situ photo of the holotype shows the proximal chevrons are more than twice as long as their corresponding centra (character 195).</span><br /><span style="background-color: #f4cccc;">- Character 205 was oddly coded inapplicable, but codes for humerofemoral length, which is 69% and thus falls into state 2.</span><br /><span style="background-color: #f4cccc;">- The deltopectoral crest apex is about 42% down the humerus (state 1), not over 50% that it was coded as in character 207.</span><br /><span style="background-color: #f4cccc;">- Like 205, <i>Chilesaurus </i>was coded inapplicable for its radiohumeral ratio, which is 70%, so state 1 in character 213.</span><br /><span style="background-color: #f4cccc;">- Since distal carpal II is unpreserved, it isn't known whether it abutted or overlapped distal carpal I (character 219).</span><br /><span style="background-color: #f4cccc;">- Similarly, its extent over metacarpal II is unknown (character 220).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was coded as nonexistent state 2 for character 221, referring to the presence of distal carpal V, which is unknown in that taxon.</span><br /><span style="background-color: #f4cccc;">- The manus is about 43% of humerus+radius length (state 1), not over 45% (state 2), for character 222.</span><br /><span style="background-color: #f4cccc;">- Metacarpal I is miscoded as being proximally narrower than metacarpal II (character 224).</span><br /><span style="background-color: #f4cccc;">- Character 232 codes for metacarpal V length, but with no character coding for metacarpal V presence, Yates coded other taxa lacking the bone as having a short mcV.&nbsp; Thus Novas et al.'s coding of inapplicable for <i>Chilesaurus </i>and <i>Tawa </i>should be changed to state 0.</span><br /><span style="background-color: #f4cccc;">- Character 242 is uncoded, though <i>Chilesaurus </i>has manual ungual II less than 70% of manual ungual I's length (state 2).</span><br /><span style="background-color: #f4cccc;">- Character 243 was coded inapplicable, but is polymorphic, as manual digit II has three phalanges (state 0) and digit III has less than four phalanges (state 1).</span><br /><span style="background-color: #f4cccc;">- Character 244 was also coded inapplicable, but should be state 1, as there are no manual digits IV or V.</span><br /><span style="background-color: #f4cccc;">- Character 252 is miscoded, as the pubic peduncle of the ilium is not twice as dorsoventrally deep as the distal end is anteroposteriorly long (state 0).</span><br /><span style="background-color: #f4cccc;">- Character 254 is also miscoded, as the ischial peduncle is not much shorter than the pubic peduncle (state 0).</span><br /><span style="background-color: #f4cccc;">- Character 255 was miscoded as having a postacetabular process longer than the distance subtended by the peduncles, when the actual value is 75% (state 0).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was coded nonexistent state 2 for character 256, as was <i>Chromogisaurus </i>(a mistake originally made by Otero and Pol).&nbsp; Both should be coded state 1, as they have a deep brevis fossa.</span><br /><span style="background-color: #f4cccc;">- Character 262 is miscoded, as <i>Chilesaurus </i>has a minimal pubic apron width of more than 40% of the width across the ilium's pubic peduncles (state 0).</span><br /><span style="background-color: #f4cccc;">- Character 267 is miscoded, as <i>Chilesaurus </i>does have a small distal pubic expansion (state 1).</span><br /><span style="background-color: #f4cccc;">- Character 278 is miscoded, as the distal ischia are narrower than deep (state 1).</span><br /><span style="background-color: #f4cccc;">- As restored, character 279 is miscoded because the hindlimb is longer than the trunk (state 0).</span><br /><span style="background-color: #f4cccc;">- Character 281 is miscoded, as the femur has a near circular section (state 0).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was coded inapplicable for anterior trochanter shape, and while it is neither a tubercle (state 0) or a ridge (state 1), it's certainly not absent (state 2).&nbsp; Thus it is coded 0/1 here.</span><br /><span style="background-color: #f4cccc;">- Character 291 is miscoded, as the anterior trochanter would be visible in posterior view (state 1).</span><br /><span style="background-color: #f4cccc;">- Another character oddly coded inapplicable, character 299 should be coded 0 (tibia longer than femur).</span><br /><span style="background-color: #f4cccc;">- Character 302 was also coded inapplicable, but should be 0 (tallest point of cnemial crest proximally located).</span><br /><span style="background-color: #f4cccc;">- Character 314 (position of iliofibularis tubercle) should be inapplicable, as the authors state <i>Chilesaurus </i>lacks one.</span><br /><span style="background-color: #f4cccc;">- There seems to be a significant fibular facet on the astragalus, making character 317 miscoded (state 0).</span><br /><span style="background-color: #f4cccc;">- Another basic limb proportion left uncoded, the metatarsotibial ratio of 62% makes character 336 coded 0.</span><br /><span style="background-color: #f4cccc;">- Character 344 seems to be miscoded, as pedal ungual I looks longer than all other non-ungual phalanges (state 1).</span><br /><span style="background-color: #f4cccc;">- Character 350 also seems to be miscoded, as pedal ungual III is less than 85% of ungual II's length (state 1).</span><br /><span style="background-color: #f4cccc;">- The final basic measurement left uncoded, the femur is between 200 and 399 mm, if the holotype's size is doubled as the author states some specimens are.</span><br /><br />The resulting tree is more resolved by <i>Chilesaurus </i>than before, with it sister to Avepoda, and successively less closely related to <i>Tawa</i>, <i>Chindesaurus </i>and <i>Agnosphitys</i>.&nbsp; Forcing it to be sauropodomorphan is now 4 steps longer instead of 5, and forcing it to be ornithischian is only 5 steps longer instead of 11.&nbsp; Thus while <i>Chilesaurus </i>had basically the same most parsimonious position, the miscodings made it seem less similar to ornithischians.&nbsp; Deleting Avepoda to test for convergence leaves <i>Chilesaurus </i>sister to <i>Tawa</i>, but now it moves to Sauropodomorpha in only one more step (Ornithischia in 4). So if <i>Chilesaurus </i>isn't an avepod, it seems basically as likely to be a sauropodomorph as a theropod, and only slightly less likely to be an ornithischian.<br /><br /><b><i>Chilesaurus </i>REtested as a basal tetanurine</b><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-IMwjOegb8vE/VYe7iYGgF9I/AAAAAAAAAoI/NFPDMInK-fg/s1600/Chile%2BAllo%2Bmanus%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="282" src="http://4.bp.blogspot.com/-IMwjOegb8vE/VYe7iYGgF9I/AAAAAAAAAoI/NFPDMInK-fg/s320/Chile%2BAllo%2Bmanus%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Chilesaurus </i>(composite using SNGM-1887, 1935 and 1937; missing apparently ultimate phalanx III-1) and <i>Allosaurus fragilis</i> (proposed neotype USNM 4734) manus scaled to same metacarpal II length (modified after Novas et al., 2015 and Gilmore, 1915 respectively).</td></tr></tbody></table><br />A fourth test involves seeing how Carrano et al.'s coelurosaur miscodings affected <i>Chilesaurus</i>' position.&nbsp; The file used is one I've been working on that recodes the included coelurosaurs (<i>Compsognathus</i>, <i>Proceratosaurus </i>and <i>Ornitholestes</i>), adds more members (<i>Bicentenaria</i>, <i>Sciurumimus</i>, <i>Zuolong</i>, <i>Juratyrant</i>, <i>Eotyrannus</i>, <i>Aorun</i>, <i>Coelurus</i>, <i>Scipionyx</i>) and corrected megaraptorans including the new <i>Megaraptor </i>skull, in addition to correcting some fragmentary taxa (<i>Poekilopleuron</i>, <i>Lourinhanosaurus</i>) and ordering a few characters correctly.&nbsp; In this test, the unstable <i>Poekilopleuron</i>, <i>Xuanhanosaurus </i>and <i>Pivetaeusaurus </i>were deleted a priori to speed up analysis time.&nbsp; None were suggested to be close to <i>Chilesaurus</i>, so it shouldn't matter.&nbsp; Again, an amazing amount of miscodings (48 of 351, or 14%) were found for <i>Chilesaurus</i>-<br /><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was left uncoded for its seemingly unfused premaxillae (character 1).</span><br /><span style="background-color: #f4cccc;">- And the moderately low premaxillary body (character 2).</span><br /><span style="background-color: #f4cccc;">- And the low angled anterior premaxillary margin (character 6).</span><br /><span style="background-color: #f4cccc;">- And the lack of a premaxillary diastema posteriorly (character 7).</span><br /><span style="background-color: #f4cccc;">- And the lack of a prominent premaxillary palatal process (character 9).</span><br /><span style="background-color: #f4cccc;">- And the seemingly non-interlocking premaxilla-maxilla articulation (character 11).</span><br /><span style="background-color: #f4cccc;">- And the at least not highly elongate anterior maxillary ramus (character 12).</span><br /><span style="background-color: #f4cccc;">- And the procumbant anterior maxillary teeth (character 13).</span><br /><span style="background-color: #f4cccc;">- And the ridge at the ventral antorbital fossa margin as coded in other matrices (character 22).</span><br /><span style="background-color: #f4cccc;">- And the lack of a supraorbital 'palpebral' (character 61).</span><br /><span style="background-color: #f4cccc;">- And the lack of postorbital-lacrimal contact (character 62).</span><br /><span style="background-color: #f4cccc;">- And the stated deep basisphenoid recess (character 96).</span><br /><span style="background-color: #f4cccc;">- There is no paradental groove due to lacking interdental plates, so character 123 is inapplicable.</span><br /><span style="background-color: #f4cccc;">- There is no lateral dentary groove, so character 124 is miscoded.</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>is left uncoded for its three premaxillary teeth (character 150).</span><br /><span style="background-color: #f4cccc;">- And its even premaxillary tooth spacing (character 151).</span><br /><span style="background-color: #f4cccc;">- And the first premaxillary tooth being subequal to the rest in size (character 152).</span><br /><span style="background-color: #f4cccc;">- And the mid-maxillary tooth spacing being adjacent (character 154).</span><br /><span style="background-color: #f4cccc;">- And the dentary teeth being subequal in size to maxillary teeth (character 155).</span><br /><span style="background-color: #f4cccc;">- Since there are no postaxial epipophyses or character coding for epipophysis presence, character 177 (epipophysis size) should be coded as the smallest state instead of inapplicable.</span><br /><span style="background-color: #f4cccc;">- Character 184 should be coded 1, as the text states <i>Chilesaurus </i>has large anterior dorsal hypapophyses.</span><br /><span style="background-color: #f4cccc;">- The cervical ribs seem to be unfused, so <i>Chilesaurus </i>should be coded for characater 208.</span><br /><span style="background-color: #f4cccc;">- Character 223 is miscoded, as the scapula is 6.9 times longer than wide (state 0).</span><br /><span style="background-color: #f4cccc;">- Character 233 is also miscoded, as the deltopectoral crest apex is ~40% down the element (state 0/1).</span><br /><span style="background-color: #f4cccc;">- The distal carpal is very comparable in shape and extent to distal carpal I of <i>Xuanhanosaurus </i>and <i>Guanlong</i>, so should be coded state 0 for character 247 (not fused to distal carpal II).</span><br /><span style="background-color: #f4cccc;">- Character264 is miscoded, as <i>Chilesaurus </i>has a narrow brevis fossa (Salgado et al., 2008).</span><br /><span style="background-color: #f4cccc;">- Character 267 is mistyped as an 8, not an actual state.&nbsp; It should be state 1 "reduced [supracetabular] shelf."</span><br /><span style="background-color: #f4cccc;">- Similarly, <i>Chilesaurus </i>has a transversely concave acetabular pubic peduncle surface (Salgado et al., 2008), so should be recoded 1 for character 269.</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>is miscoded as having a pubic peduncle much larger than the ischial peduncle (character 270)</span><br /><span style="background-color: #f4cccc;">- Character 271 is miscoded, as <i>Chilesaurus </i>does not have an acuminate ischial peduncle.</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>has a pubic peduncle over twice as long as wide distally (Salgado et al., 2008),so was miscoded for character 272.</span><br /><span style="background-color: #f4cccc;">- Character 276 is also miscoded, as <i>Chilesaurus </i>has a ventral preacetabular lobe (state 1).</span><br /><span style="background-color: #f4cccc;">- Character 281 is miscoded, as <i>Chilesaurus </i>lacks a fenestra in its pubis.</span><br /><span style="background-color: #f4cccc;">- Character 284 is miscoded as well, as the pubic apices don't contact (state 0).</span><br /><span style="background-color: #f4cccc;">- As there is no obturator foramen, character 287 should be coded 0.</span><br /><span style="background-color: #f4cccc;">- While the authors code <i>Chilesaurus </i>as inapplicable for character 290 "Pubis, shape of boot in ventral view", Carrano et al. code taxa with no substantial boot based on their distal proportion.&nbsp; So given <i>Chilesaurus</i>' wide distal end, it should be coded 0 as well.</span><br /><span style="background-color: #f4cccc;">- Character 297 is miscoded, as <i>Chilesaurus </i>has an ischial apron. </span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>lacks a dorsomedian ischial ridge (character 298).</span><br /><span style="background-color: #f4cccc;">- It is miscoded for character 307, as <i>Chilesaurus </i>has a prominent fourth trochanter.</span><br /><span style="background-color: #f4cccc;">- Similar to character 290, as <i>Chilesaurus </i>lacks a mediodistal femoral crest, it should be coded as the less developed state of character 310, not inapplicable.</span><br /><span style="background-color: #f4cccc;">- Characters 323 and 324 are miscoded, as <i>Chilesaurus </i>lacks a fibular crest so would be inapplicable for thickness and connection to lateral condyle.</span><br /><span style="background-color: #f4cccc;">- In the same way, the fibula is stated to lack a proximomedial fossa, so should be coded inapplicable for character 325 and 326 describing its morphology.</span><br /><span style="background-color: #f4cccc;">- The authors state <i>Chilesaurus </i>lacks an iliofibularis tubercle on the fibula, so it should be coded 0 for character 327, not unknown.</span><br /><span style="background-color: #f4cccc;">-&nbsp; The authors didn't notice the mistake in Carrano et al.'s matrix where character 331 ("Astragalus, ascending process morphology: absent (0), blocky (1), laminar (2).") is coded with only two states, blocky as 0 and laminar as 1.&nbsp; Thus <i>Chilesaurus</i>' 1 coding should be 0.</span><br /><span style="background-color: #f4cccc;">- Character 333 is miscoded, as the astragalar ascending process is lower than the astragalar body (state 0).</span><br /><span style="background-color: #f4cccc;">- Character 335 is also miscoded, as the astragalar fossa in <i>Chilesaurus </i>is posterior to the ascending process, so it has no fossa anterior to the process (0).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>seems to have a wide fibular fossa on its astragalus, so would be recoded 0 for character 336.</span><br /><span style="background-color: #f4cccc;">- Metatarsal I is over 50% of metatarsal II's length (character 340).</span><br /><br />Once these rather numerous miscodings and uncodings are corrected for, <i>Chilesaurus </i>emerges as the most basal megalosaurid.&nbsp; As it only took two steps to place in Megalosauroidea before the corrections, this isn't much of a change.&nbsp; What IS a big change though is that it now takes only two more steps to place within Coelurosauria (in a clade with <i>Aorun</i> and <i>Sciurumimus</i>) or outside Avepoda.&nbsp; Before the corrections, the number of steps needed was 7 and 13 respectively, so the miscodings heavily weighted it toward the basal tetanurine position.&nbsp; This shows <i>Chilesaurus </i>isn't strongly supported as a basal tetanurine.<br /><br />Note I didn't bother recoding <i>Chilesaurus </i>in Smith et al.'s matrix because my prior work on it (Mortimer and Marjanovic, in prep.) indicates there are large amounts of uncoded characters for many taxa.&nbsp; But recall from the last post that only two steps were needed to move <i>Chilesaurus </i>outside Avepoda, and that at least two miscodings were found that would do that. &nbsp; <br /><br /><b><i>Chilesaurus </i>REtested as a dinosaur </b><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-nKpAalAuuG4/VYfDDvbrhtI/AAAAAAAAAoc/IWUG2zoAflU/s1600/Chile%2BEust%2Bastragalus%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="113" src="http://3.bp.blogspot.com/-nKpAalAuuG4/VYfDDvbrhtI/AAAAAAAAAoc/IWUG2zoAflU/s320/Chile%2BEust%2Bastragalus%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Chilesaurus </i>(paratype SNGM-1936) and <i>Eustreptospondylus </i>(holotype OUM J13558) astragali in anterior view scaled to same width (modified after Novas et al., 2015 and Sadlier et al., 2008 respectively).</td></tr></tbody></table><br />My fifth and final test used Nesbitt et al.'s matrix, but with <i>Daemonosaurus </i>added as in Sues et al. (2011) and changes from Langer and Ferigolo (2013).&nbsp; I also added the characters from Nesbitt (2011) that vary in Dinosauromorpha (22, 58, 85, 113, 127, 129, 255), <i>Lutungutali</i> (codings from Peecook et al., 2013), <i>Pseudolagosuchus</i> (codings from Nesbitt, 2011), <i>Lewisuchus</i> (codings from Bittencourt et al., 2014), <i>Asilisaurus</i> (codings from Nesbitt, 2011), <i>Diodorus</i> (codings from Kammerer et al., 2012), <i>Panphagia</i>, <i>Pampadromaeus</i> (codings partly from Cabreira et al., 2011), <i>Chromogisaurus</i>, <i>Agnosphitys</i>, <i>Guaibasaurus</i> (codings from Ezcurra, 2010), <i>Saltopus</i> (codings from Langer and Ferigolo, 2013), <i>Teyuwasu</i>, <i>Nyasasaurus</i> (codings from Nesbitt et al., 2013), <i>Eodromeus</i> and the new characters from the <i>Daemonosaurus</i>, <i>Lutungutali, Diodorus</i>, <i>Nyasasaurus </i>and <i>Guaibasaurus </i>(Ezcurra, 2010) analyses.&nbsp; <i>Velociraptor </i>(75 miscodings) and <i>Eoraptor </i>(91 miscodings) were comprehensibly checked for accuracy, and I ended up finding a lot of other theropod miscodings too. In the case of <i>Chilesaurus</i>, 38 of 315 or 12% of characters were miscoded.<br /><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was left uncoded for character 1, the proportions of the subnarial premaxilla, which are state 1.</span><br /><span style="background-color: #f4cccc;">- Based on the anterior body's slope, the premaxilla's dorsal process would have state 1 for character 3 instead of ?.</span><br /><span style="background-color: #f4cccc;">- The premaxilla has a long subnarial process, so is state 1 for character 4 instead of ?.</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was miscoded as unknown for character 5, when the authors state its premaxilla has a "prominent plate-like postnarial process."</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>was left uncoded for the absence of a posteroventral premaxillary process (character 7, state 0). </span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>is left uncoded for premaxillary tooth number (character 9), which was coded as being 3 in the Smith et al. matrix.</span><br /><span style="background-color: #f4cccc;">- It was left uncoded for character 11, though it clearly has state 0 of premaxillary teeth present anteriorly.</span><br /><span style="background-color: #f4cccc;">- Character 50 was left uncoded, but the skeletal indicates <i>Chilesaurus </i>lacks pseudosuchian-like ventrally extensive postorbital-squamosal contact.</span><br /><span style="background-color: #f4cccc;">- The authors stare <i>Chilesaurus </i>has a deep basisphenoid recess, so should be coded 1 for character 69, not ?.</span><br /><span style="background-color: #f4cccc;">- Character 85 should be a no-brainer to code (skull less than 50% of presacral length), but was inexcusably left uncoded. </span><br /><span style="background-color: #f4cccc;">- Again, the orbit is clearly round based on the frontal, so character 89 should be coded.</span><br /><span style="background-color: #f4cccc;">- Character 111 is miscoded, as <i>Chilesaurus </i>lacks "apicobasally tall and blade-like" teeth.</span><br /><span style="background-color: #f4cccc;">- The figured cervical 4 seems to lack epipophyses, and it's coded as having"absent or poorly developed" postaxial epipophyses in the Smith et al. matrix and none in the Yates matrix, which would make character 126 miscoded.</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>has a forelimb/hindlimb ratio of 58%, so should be coded 0 for character 150.</span><br /><span style="background-color: #f4cccc;">- The coracoid is stated to lack a biceps tuber by the authors, contra their coding of character 158.</span><br /><span style="background-color: #f4cccc;">- 161 should be coded 0 (humerofemoral ratio over 59%) , as <i>Chilesaurus</i>' ratio is 69%.</span><br /><span style="background-color: #f4cccc;">- The humerus has a head separated from the deltopectoral crest by a narrow ridge, so <i>Chilesaurus </i>should be coded 1 for character 163.</span><br /><span style="background-color: #f4cccc;">- The metacarpometatarsal ratio is .37 based on the measurement table and manus figure, so should be coded 1 for character 172, not 0.</span><br /><span style="background-color: #f4cccc;">- Character 179 should be coded 0 ("Digit I with metacarpal: longer than the ungual") instead of ?.</span><br /><span style="background-color: #f4cccc;">- Character 188 should be coded 2, as manual digit V is absent or "reduced to a tiny nubbin".&nbsp; This showed nothing was coded as state 2, and indeed <i>Allosaurus</i>, <i>Velociraptor</i>, <i>Ceratosaurus</i>, <i>Dilophosaurus</i>, <i>Coelophysis</i>, <i>Tawa</i>, <i>Eudimorphodon</i> and <i>Dimorphodon </i>should be coded that way too.</span><br /><span style="background-color: #f4cccc;">- As <i>Chilesaurus </i>lacks a supracetabular crest, it cannot be coded for that crest's orientation (character 189).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>has a basically squared anterior preacetabular margin, so should be coded 1 for character 193, not ?.</span><br /><span style="background-color: #f4cccc;">- Similarly, character 194 should be coded 0 ("posterior margin of the postacetabular process in lateral view: straight or</span><br /><span style="background-color: #f4cccc;">convex"), not ?.</span><br /><span style="background-color: #f4cccc;">- Character 202 is miscoded, as <i>Chilesaurus </i>doesn't have a posteriorly hooked ischial peduncle.</span><br /><span style="background-color: #f4cccc;">- Character 203 is also miscoded according to the measurement table, as <i>Chilesaurus </i>has a pubofemoral ratio of less than 70%.</span><br /><span style="background-color: #f4cccc;">- In a miscoding that makes <i>Chilesaurus </i>actually less like ornithischians, it doesn't seem to have a prepubis (character 205).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>is miscoded as lacking a pubic boot (character 207). </span><br /><span style="background-color: #f4cccc;">- The pubic boot is less than 33% of pubic length (character 209).</span><br /><span style="background-color: #f4cccc;">- Character 210 is miscoded, as the figure clearly shows the pubic articular surfaces for ilium and ischium are separated by a gap.</span><br /><span style="background-color: #f4cccc;">- The pubes are distally narrower than proximally, so character 213 should be recoded 1. </span><br /><span style="background-color: #f4cccc;">- In the same way, character 219 should be coded 2 ("Ischium, proximal articular surfaces: articular surfaces with the ilium and the pubis separated by a large concave surface"), not 0.</span><br /><span style="background-color: #f4cccc;">- Based on the prominent anterior trochanter, it's near certain <i>Chilesaurus </i>was miscoded as having a silesaur-like flat "anterolateral" proximal femoral surface (character 225).</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>lacks a dorsolateral femoral trochanter (state 0), so is miscoded for character 230. </span><br /><span style="background-color: #f4cccc;">- Character 265 is miscoded, as the authors state the proximomedial fibular face has no concavity </span><br /><span style="background-color: #f4cccc;">- Character 273 is miscoded, as <i>Chilesaurus </i>has an astragalar ascending process shorter than the astragalar body.</span><br /><span style="background-color: #f4cccc;">- Similarly, the ascending process is blocky, not laminar.&nbsp; So 274 is miscoded.</span><br /><span style="background-color: #f4cccc;">- <i>Chilesaurus </i>lacks osteoderms, so should be coded 0 for character 308.</span><br /><span style="background-color: #f4cccc;">- Finally, <i>Chilesaurus </i>is coded in the Smith et al. matrix as lacking gastralia, so should be coded 2 for character 315.</span><br /><br /><i>Chilesaurus </i>still emerges as the sister group of the only included coelurosaur, <i>Velociraptor</i>.&nbsp; But now that both are coded correctly, <i>Chilesaurus </i>needs 9 more steps to group with basal tetanurines. So the matrix doesn't just generically support a neotheropod assignment for <i>Chilesaurus</i>, it only appeared to because <i>Velociraptor </i>was wrongly coded the same as <i>Allosaurus </i>and <i>Ceratosaurus </i>for so many characters.&nbsp; Forcing <i>Chilesaurus </i>outside of Avepoda takes 11 more steps, and it becomes the sister group of Avepoda, though the clade reverses its polarity so that <i>Velociraptor </i>is the first to branch off and coelophysids are 'most derived'.&nbsp; Forcing it to either Sauropodomorpha or Ornithischia takes 17 more steps.&nbsp; These sound like easily rejected alternatives, except that most of the steps are needed to get <i>Chilesaurus </i>away from <i>Velociraptor </i>itself, not Neotheropoda as a whole.&nbsp; We can tell this by deleting <i>Velociraptor </i>and rerunning things.&nbsp; <i>Chilesaurus </i>remains closest to neotheropods, but now it only takes 7 more steps to place it in Sauropodomorpha or Ornithischia.&nbsp; Furthermore, if we force it out of Avepoda, it takes one of these positions instead of still being a theropod, since the artificially 'basal' <i>Velociraptor </i>isn't there to attract it.<br /><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">|--<i>Marasuchus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">`--+--<i>Teyuwasu</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp;&nbsp; `--Dinosauria</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; |--Ornithischia</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; |&nbsp; |--<i>Lewisuchus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; |&nbsp; `--+--+--<i>Pseudolagosuchus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; |&nbsp; `--<i>Asilisaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; `--+--Silesauridae</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp; |--<i>Eucoelophysis</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--+--+--<i>Lutungutali</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; |&nbsp; `--<i>Silesaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; `--+--<i>Diodorus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--<i>Sacisaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--Genasauria</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--<i>Scutellosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--<i>Eocursor</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--<i>Lesothosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--<i>Pisanosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--<i>Heterodontosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; `--Saurischia</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |*-<i>Saltopus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |--Herrerasauridae</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |--<i>Staurikosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--+--<i>Chindesaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; `--<i>Herrerasaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--Eusaurischia</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--Sauropodomorpha</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; |--<i>Guaibasaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |&nbsp; `--+--+--<i>Efraasia</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; |&nbsp; `--<i>Plateosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; `--+--+--<i>Panphagia</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--<i>Pampadromaeus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp;&nbsp; `--Saturnaliinae</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--<i>Chromogisaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; | &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--<i>Saturnalia</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--Theropoda</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |--<i>Agnosphitys</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--<i>Eoraptor</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--<i>Nyasasaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--<i>Eodromaeus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--+--<i><b>Chilesaurus</b></i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--<i>Daemonosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--<i>Tawa</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--Avepoda</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp; |--Coelophysoidea</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; |--<i>Liliensternus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |&nbsp; `--+--<i>kayentakatae</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; | &nbsp; &nbsp; `--<i>Coelophysis</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--<i>Zupaysaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--+--<i>Cryolophosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--+--<i>Dilophosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--Neotheropoda</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; |--<i>Ceratosaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp; `--Tetanurae</span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; |--<i>Piatnitzkysaurus</i></span><br /><span style="font-family: &quot;Courier New&quot;, Courier, monospace;">&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; `--<i>Allosaurus</i></span><br /><br /><div style="text-align: center;"><span style="font-family: &quot;Courier New&quot;, Courier, monospace;"><span style="font-family: inherit;"><span style="font-family: Times,&quot;Times New Roman&quot;,serif;"><span style="font-family: inherit;">One m</span>ost parsimonious tree of modified Nesbitt et al. matrix after deletion of <i>Saltopus </i>and <i>Velociraptor</i>, using only verifiable codings for <i>Chilesaurus </i>(see below).&nbsp;</span></span> </span></div><br /><b>So many miscodings</b> <br /><br />I gotta say that I'm surprised each matrix has at least 12-14% of <i>Chilesaurus</i>' characters miscoded.&nbsp; That's likely to be an underestimate since the taxon was described in a tabloid (Nature), so e.g. the vertebrae are almost completely undescribed and unillustrated.&nbsp; Indeed, some of the coded characters that cannot be checked would be very odd for a dinosaur if true.&nbsp; For instance, <i>Chilesaurus </i>is coded as having a hooked metatarsal V in the Nesbitt et al. matrix.&nbsp; Yet this isn't found in any known dinosauromorph.&nbsp; This makes me wonder if <i>Chilesaurus </i>is falling out in so many different positions because its actual morphology isn't being tested.&nbsp; So I recoded <i>Chilesaurus </i>for only those characters that could be determined from descriptions or illustrations, not merely the authors' codings.<br /><br />In the Carrano et al. matrix, verifiable <i>Chilesaurus </i>emerges as the most basal tetanurine, or just closer to birds than <i>Chuandongocoelurus </i>and <i>sinensis</i>, or in Coelurosauria (as the sister of <i>Aorun</i>).&nbsp; Only a single extra step is needed place it outside Avepoda now.<br /><br />In the Otero and Pol matrix, verifiable <i>Chilesaurus </i>emerges in a trichotomy with <i>Tawa </i>and Avepoda. &nbsp; It now only takes two more steps to place in Sauropodomorpha (emerges as the basalmost), and four more in Ornithischia (down from 4 and 5 respectively).&nbsp; <br /><br />In the Nesbitt et al. matrix, verifiable <i>Chilesaurus </i>still emerges sister to <i>Velociraptor</i>.&nbsp; Now 9 steps are needed to exclude it from Avepoda (emerges as sister to <i>Daemonosaurus</i>), but these are all due to <i>Velociraptor</i>.&nbsp; For if <i>Velociraptor </i>is deleted while using verifiable <i>Chilesaurus</i>, the latter emerges as either sister to the two tetanurines or sister to <i>Daemonosaurus </i>outside of Avepoda.&nbsp; Using verifiable <i>Chilesaurus </i>really helps it group with ornithischians, taking only 7 more steps instead of 17, which doesn't change much (8 more steps vs. 7) if <i>Velociraptor </i>is deleted.&nbsp; This shows the steps that grouped <i>Chilesaurus </i>with <i>Velociraptor </i>to the exclusion of ornithischians were largely those that cannot be verified in <i>Chilesaurus </i>given its description.&nbsp; On the other hand, using only verifiable <i>Chilesaurus </i>characters made it slightly less likely to be sauropodomorphan (23 more steps vs. 17), but if <i>Velociraptor </i>is deleted this falls to 7 more steps just like if even the questionable <i>Chilesaurus </i>characters are retained.<br /><br /><b>What's it mean? </b><br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://1.bp.blogspot.com/-FeSTX5iqKu0/VYfNLDB5jOI/AAAAAAAAAow/Jnao8qZiwqg/s1600/Chile%2Bskel%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="171" src="http://1.bp.blogspot.com/-FeSTX5iqKu0/VYfNLDB5jOI/AAAAAAAAAow/Jnao8qZiwqg/s320/Chile%2Bskel%2Bcomp.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Chilesaurus (after Novas et al., 2015; lower left) compared to its suggested relatives- Monolophosaurus (copyright Scott Hartman; upper left), Shuvuuia (copyright Ville Sinkkonen; upper right), and Tawa (copyright Scott Hartman; lower right).&nbsp; Scaled to same femoral length.</td></tr></tbody></table><br /><br />My conclusion here (finally!) is that using pure parsimony, <i>Chilesaurus </i>will clade with maniraptoriforms and more precisely alvarezsaurids.&nbsp; It emerged strongly supported in that position in the Lori matrix and Nesbitt et al.'s once only verified codings were used, took only 4 steps to move there without any theropod characters in Butler's ornithischian matrix, and is one of the most parsimonious possibilities in Carrano et al.'s matrix if only its verifiable codings are used.&nbsp; A basal tetanurine position is strongly rejected if alvarezsaurids are included (13 more steps in the Lori matrix; 10 more in Butler's matrix). Yet I think both of these placements are unlikely due to the incongruities caused by inserting <i>Chilesaurus </i>there, so that even though it's not most parsimonious, I find convergence between <i>Chilesaurus </i>and tetanurines more likely than reversals in <i>Chilesaurus</i>.&nbsp; The fact verifiable <i>Chilesaurus </i>emerges outside Avepoda in one of the most parsimonious trees when <i>Velociraptor </i>is excluded from Nesbitt et al.'s matrix and only takes 1 more step to place there in Carrano et al.'s matrix thus makes sense to me.&nbsp; This would allow many characters to be plesiomorphic (e.g. no fibular crest, short astragalar ascending process, large pedal digit I), and also agrees with its placement in Otero and Pol's trees.&nbsp; I admit I might be proven wrong if this is a repeat of 1984-1992 Segnosauria and we find a proto-chilesaur with more megalosauroid or alvarezsaurid characters, but for now this seems more realistic to me (see my preferred cladogram above).&nbsp; Is it possibly not theropod?&nbsp; Verifiable <i>Chilesaurus </i>falls out in Sauropodomorpha with 2-7 more steps, and in Ornithischia with 4-8 more.&nbsp; So it seems possible it's either one, and combining the Yates and Nesbitt matrix would give us better numbers.&nbsp; But what we really need before a much better conclusion can be reached is an <b>osteology of <i>Chilesaurus</i></b>.&nbsp; Please say one's in development.&nbsp; The thing's simply too weird, with too many coding issues for a tabloid description to work long term.&nbsp; <br /><br /><b>References</b>- Gilmore, 1915. On the fore limb of <i>Allosaurus fragilis</i>. Proceedings of the United States National Museum. 49, 501-513.<br /><br />Welles, 1984. <i>Dilophosaurus wetherilli</i> (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.<br /><br />Bonaparte, 1986. Les Dinosaures (Carnosaures, Allosauridés, Sauropodes, Cétiosauridés) du Jurassique moyen de Cerro Cóndor (Chubut, Argentine). Annales de Paléontologie. 72, 247-289.<br /><br />Zhao and Currie, 1994. A large crested theropod from the Jurassic of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences. 30(10), 2027-2036. <br /><br />Sadlier, Barrett and Powell, 2008. The anatomy and systematics of <i>Eustreptospondylus oxoniensis</i>, a theropod dinosaur from the Middle Jurassic from Oxfordshire, England. Monograph of the Palaeontological Society. 1-82.<br /><br />Salgado, Cruz, Suarez, Fernandez, Gasparini, Palma-Heldt and Fanning, 2008. First Late Jurassic dinosaur bones from Chile. Journal of Vertebrate Paleontology. 28(2), 529-534.<br /><br />Alifanov and Barsbold, 2009. <i>Ceratonykus oculatus</i> gen. et sp. nov., a new dinosaur (?Theropoda, Alvarezsauria) from the Late Cretaceous of Mongolia. Paleontological Journal (English edition). 43(1), 94-106.<br /><br />Brusatte, Benson, Currie and Zhao, 2010. The skull of <i>Monolophosaurus jiangi</i> (Dinosauria: Theropoda) and its implications for early theropod phylogeny and evolution. Zoological Journal of the Linnean Society. 158(3), 573-607. <br /><br />Ezcurra, 2010. A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: A reassessment of dinosaur origin and phylogeny. Journal of Systematic Palaeontology. 8(3), 371-425.&nbsp; <br /><br />Alifanov and Saveliev, 2011. Brain structure and neurobiology of alvarezsaurians (Dinosauria), exemplified by <i>Ceratonykus oculatus</i> (Parvicursoridae) from the Late Cretaceous of Mongolia. Paleontological Journal. 45(2), 183-190.<br /><br />Cabreira, Schultz, Bittencourt, Soares, Fortier, Silva and Langer, 2011. New stem-sauropodomorph (Dinosauria, Saurischia) from the Triassic of Brazil. Naturwissenschaften. 98(12), 1035-1040. <br /><br />Nesbitt, 2011. The early evolution of archosaurs: Relationships and the origin of major clades. 352, 292 pp.<br /><br />Sues, Nesbitt, Berman and Henrici, 2011. A late-surviving basal theropod dinosaur from the latest Triassic of North America. Proceedings of the Royal Society B. 278(1723), 3459-3464. <br /><br />Han, Barrett, Butler and Xu, 2012. Postcranial anatomy of <i>Jeholosaurus shangyuanensis</i> (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China. Journal of Vertebrate Paleontology. 32(6), 1370-1395.<br /><br /><span class="reference-text">Kammerer, Nesbitt and Shubin, 2012. The first basal dinosauriform (Silesauridae) from the Late Triassic of Morocco. Acta Palaeontologica Polonica. 57(2), 277-284</span>.<br /><br />Langer and Ferigolo, 2013. The Late Triassic dinosauromorph <i>Sacisaurus agudoensis</i> (Caturrita Formation; Rio Grande do Sul, Brazil): Anatomy and affinities. In Nesbitt, Desojo and Irmis (eds.). Anatomy, Phylogeny and Palaeobiology of Early<br />Archosaurs and their Kin. Geological Society, London, Special Publicataions. 379, 353-392.<br /><br />Nesbitt, Barrett, Werning, Sidor and Charig, 2013. The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania. Biology Letters. 9(1), 20120949. <br /><br />Peecook, Sidor, Nesbitt, Smith, Steyer and Angielczyk, 2013. A new silesaurid from the upper Ntawere Formation of Zambia (Middle Triassic) demonstrates the rapid diversification of Silesauridae (Avemetatarsalia, Dinosauriformes). Journal of Vertebrate Paleontology. 33(5), 1127-1137.<br /><br />Bittencourt, Arcucci, Marsicano and Langer, 2014. Osteology of the Middle Triassic archosaur <i>Lewisuchus admixtus</i> Romer (Chañares Formation, Argentina), its inclusivity, and relationships amongst early dinosauromorphs. Journal of Systematic Palaeontology. 13(3), 189-219.<br /><br />Godefroit, Sinitsa, Dhouailly, Bolotsky, Sizov, McNamara, Benton and Spagna, 2014. A Jurassic ornithischian dinosaur from<br />Siberia with both feathers and scales. Science. 345(6195), 451-455.<br /><br />Novas, Salgado, Suarez, Agnolın, Ezcurra, Chimento, Cruz, Isasi, Vargas and Rubilar-Rogers, 2015. An enigmatic plant-eating theropod from the Late Jurassic period of Chile. Nature. 522, 331-334. <br /><br /><!--70--><!--50-->Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com1tag:blogger.com,1999:blog-3248412803814730250.post-2017650452213030792015-05-20T05:25:00.000-07:002015-05-20T05:25:38.535-07:00Is Chilesaurus a basal tetanurine?I've been away for a few weeks, but came back to find the weird new dinosaur <i>Chilesaurus </i>exists.&nbsp; Just looking at it, any dinosaur worker would notice an odd mix of characters, making me curious exactly what it's related to.&nbsp; Novas et al. (2015) used four analyses to determine it belonged in non-orionidan Tetanurae.&nbsp; To use a Peters-ism, that's one strange bed fellow.&nbsp; So let's go in for a closer look.<br /><br /><b>Their analyses </b><br /><br />The first analysis was Nesbitt et al.'s (2009) <i>Tawa </i>analysis (315 characters), which focused on basal theropods and saurischians.&nbsp; This found <i>Chilesaurus </i>to be a coelurosaur, though so was <i>Ceratosaurus</i>.&nbsp; The authors rightly note the matrix was not designed to test neotheropod phylogeny, and the low Bremer supports show this, though I note the pairing of <i>Chilesaurus </i>with the only included actual coelurosaur <i>Velociraptor </i>is better supported than the other neotheropodan nodes. 15 more steps were needed to constrain it as sister to Avepoda, 16 more to constrain it as a sauropodomorph (ended up sister to <i>Plateosaurus </i>instead of <i>Saturnalia </i>or <i>Efraasia</i>), and 14 more as an ornithischian (as the basalmost one or a heterodontosaurid).&nbsp; These sound impressive until you realize that the matrix wasn't designed with ornithischian or sauropodomorph apomorphies in mind.&nbsp; Also, I wonder how many more steps it took to place as a basal tetanurine where Novas et al. ultimately conclude it goes?&nbsp; And why not use Nesbitt's (2011) larger anlysis which expanded on this and has had a major update by Langer and Ferigolo (2013), or at least Sues et al.'s (2011) version of Nesbitt et al.'s that added <i>Daemonosaurus</i>?&nbsp; The latter genus also has three premaxillary teeth, a short snout, broad posterolateral premaxillary process and elongate cervicals with pleurocoels, so might help move <i>Chilesaurus </i>out of Avepoda.<br /><br />The second analysis is a logical follow up, coding <i>Chilesaurus </i>in the sauropodomorph-focused matrix of Otero and Pol (2013) (with <i>Tawa </i>added; 353 characters), that is itself basically Yates' 2007 'prosauropod' matrix.&nbsp; This finds <i>Chilesaurus </i>in a polytomy with Avepoda, <i>Tawa </i>and <i>Chindesaurus</i>, but placing it in Sauropodomorpha (as the most basal member) is now only five steps longer.&nbsp; I'd say that's not strong support for <i>Chilesaurus </i>being a theropod, as numerous relationships rejected by five steps in one matrix end up being supported in other or later matrices.&nbsp; It now takes 11 more steps to place it sister to Ornithischia, but that OTU isn't divided up, making the situation more problematic than above.<br /><br />The third analysis uses a revised version of Smith et al.'s (2007) theropod matrix <a href="http://theropoddatabase.blogspot.com/2010/12/adventures-in-not-coding-marasuchus-by.html">that is terribly mis/un-coded</a>.&nbsp; Not encouraging, though Novas et al. "deeply rescored" "several taxa" "based on new available data" including at least <i>Eoraptor</i> and <i>Megaraptor</i>.&nbsp; They also added <i>Tawa</i>, <i>Aerosteon</i>, <i>Falcarius</i>, <i>Jianchangosaurus </i>and Therizinosauridae and 56 new characters (total now 412 characters).&nbsp; Here, <i>Chilesaurus </i>emerges as sister to Piatnitzkysauridae+Orionides.&nbsp; However, only TWO steps are needed to constrain it as the sister of Avepoda*, and only three are needed to place it as the basalmost coelurosaur.&nbsp; So even if the authors fixed the matrix, it only very weakly supports a basal tetanurine position.&nbsp; Suspiciously, <i>Plateosaurus </i>was deleted from the matrix "because of the morphological gap present between the very early sauropodomorphs present in the data set (e.g. <i>Saturnalia</i>) and <i>Plateosaurus</i>."&nbsp; This is rich when they left <i>Velociraptor </i>in Nesbitt et al.'s matrix, which is at least as different from <i>Allosaurus</i>.&nbsp; As the only sauropodomorphs left were the incomplete <i>Saturnalia </i>and controversial <i>Eoraptor</i>, and considering <i>Chilesaurus </i>emerged sister to <i>Plateosaurus </i>when constrained as a sauropodomorph in their first analysis, I can't help but wonder if <i>Chilesaurus </i>was sister to <i>Plateosaurus </i>here too until the latter was deleted.&nbsp; Novas et al. never say how many more steps it takes to place <i>Chilesaurus </i>in Sauropodomorpha in this matrix, and ornithischians weren't included.<br /><br />* Just looking through their list of characters supporting the position, astragalar ascending process height is miscoded in Smith et al.'s (it's clearly not "higher than the astragalar body, typically covering only lateral half of anterior surface of distal tibia"), as is "ridge on lateral side of tibia for connection with fibula present and clearly separated from proximal articular surface."&nbsp; So there's the two steps we need to move it outside Avepoda. <br /><br />Finally, the fourth analysis used Carrano et al.'s (2012) basal tetanurine-focused analysis (351 characters).&nbsp; Now <i>Chilesaurus</i> emerges in a polytomy with <i>Monolophosaurus</i>, <i>Chuandongocoelurus</i>, Megalosauroidea (including piatnitzkysaurids) and Avetheropoda.&nbsp; Again, only two more steps are necessary to place it in Megalosauroidea, so its precise position here is very poorly supported.&nbsp; Seven more steps are needed to place it in Coelurosauria (emerges sister to <i>Compsognathus</i>), but only three coelurosaurs are included in the matrix and they're heavily miscoded.&nbsp; I added a lot of basal coelurosaurs to it when testing <i>Bahariasaurus </i>and found e.g. <i>Compsognathus </i>has <u><b>112 miscodings</b></u> (of 351 characters).&nbsp; That these include at least 7 that are actually more similar to <i>Chilesaurus </i>seems likely, though of course they probably also include at least 7 less similar to it, with the conclusion that the original matrix can't tell us how <i>Chilesaurus </i>compares to coelurosaurs.&nbsp; As no maniraptoriforms were included, it can't tell us if <i> Chilesaurus </i>might belong there either.&nbsp; No sauropodomorphs except the controversial <i>Eoraptor </i>were included, and no ornithischians.&nbsp; Though like above matrix, this one wasn't made to test such basal nodes.<br /><br />Given these results, Novas et al.'s conclusion seems to be stated far more strongly than their evidence indicates.&nbsp; They say "the four independent phylogenetic data matrix [sic] favours a position as a neotheropod [avepod in my terminology], and particularly as a basal tetanuran", "The results of the four analyses are detailed below, but all of them agree in the position of <i>Chilesaurus </i>as a tetanuran theropod", and "Remarkably, all these analyses placed <i>Chilesaurus </i>as a member of Theropoda, near the origin of tetanurans."&nbsp; Only the first two matrices have the needed basal taxa to test whether <i>Chilesaurus </i>is in Avepoda, and the second doesn't place it any closer to avepods than <i>Tawa </i>or <i>Chindesaurus </i>and supports Sauropodomorpha as a highly plausible alternative.&nbsp; Further, the first places it as a ceratosaur not a tetanurine, the second doesn't even split Avepoda into multiple OTUs so can't weigh in, and the third is basically ambiguous whether <i>Chilesaurus </i>is actually sister to Avepoda or a coelurosaur.&nbsp; Only the fourth analysis may strongly support placing <i>Chilesaurus </i>as a monolophosaur-piatnitzkysaur grade tetanurine, though we don't know how easy it is to place outside Avepoda in that one.<br /><br />I'd say that even if we trusted these matrices accuracy 100%, they only tell us it's quite possible <i>Chilesaurus </i>is the basalmost sauropodomorph or coelurosaur and is about equally likely to be sister to Avepoda, a non-orionidan tetanurine or a basal megalosauroid.&nbsp; Not actually much agreement there.<br /><br /><b>My analyses</b><br /><br />Will come next... <br /><br /><b>References</b>- Smith, Makovicky, Hammer and Currie, 2007. Osteology of <i>Cryolophosaurus ellioti</i> (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.<br /><br />Yates, 2007. The first complete skull of the Triassic dinosaur <i>Melanorosaurus </i>Haughton (Sauropodomorpha, Anchisauria). Special Papers in Palaeontology. 77, 9-55.<br /><br />Nesbitt, Smith, Irmis, Turner, Downs and Norell, 2009. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science. 326, 1530-1533.<br /><br />Nesbitt, 2011. <span class="st">The early evolution of archosaurs: Relationships and the origin of major clades. Bulletin of the American Museum of Natural History. 352, 292 pp.</span><br /><br /><span class="st">Sues, Nesbitt, Berman and Henrici, 2011. A late-surviving basal theropod dinosaur from the latest Triassic of North America. Proceedings of the Royal Society B. 278(1723), 3459-3464.</span><br /><br /><span class="st">Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300. </span><br /><br /><span class="st">Langer and Ferigolo, 2013. The Late Triassic dinosauromorph <i>Sacisaurus agudoensis</i> (Caturrita Formation; Rio Grande do Sul, Brazil): Anatomy and affinities. </span>Geological Society, London, Special Publications. 379(1), 353-392.<br /><br />Otero and Pol, 2013. Postcranial anatomy and phylogenetic relationships of <i>Mussaurus patagonicus</i> (Dinosauria, Sauropodomorpha). Journal of Vertebrate Paleontology. 33(5), 1138-1168.<br /><br />Novas, Salgado, Suarez, Agnolın, Ezcurra, Chimento, Cruz, Isasi, Vargas and Rubilar-Rogers, 2015. An enigmatic plant-eating theropod from the Late Jurassic period of Chile. Nature. doi:10.1038/nature14307Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com1tag:blogger.com,1999:blog-3248412803814730250.post-10979367681945564592015-02-26T05:27:00.002-08:002015-02-26T05:27:47.965-08:00Lingham-Soliar's (2015) Galileo complexHooo-eeee, Theagarten Lingham-Soliar (2015) just published volume 2 of his work "The Vertebrate Integument."&nbsp; Sure sounds scholarly, and parts of it are, but the last chapter is something else.<br /><br /><b>Before the end </b><br /><br />The preceding chapter also has some fun parts, such as section 6.4- "<i>Sinosauropteryx</i>, A Basal Theropod."&nbsp; Well, it's not exactly basal, that would be more like <i>Eodromaeus</i>, or coelophysoids.&nbsp; He then goes on to say "Given the importance of <i>Sinosauropteryx </i>as a basal dinosaur with alleged protofeathers."&nbsp; Wait, now it's a basal dinosaur?!&nbsp; He repeats this again in section 7.4.1.&nbsp; Try 'basal coelurosaur' if you want the term to be meaningful.<br /><br />Lingham-Soliar follows this with section 6.5- "Pushing Feathered Dinosaurs into the Mid-Triassic", based on Xu et al.'s (2009) <i>Beipiaosaurus </i>paper.&nbsp; The Mid-Triassic part is based on a quote in their concluding paragraph where they note the resemblance of coelurosaurian integument to <i>Psittacosaurus</i>' quills and pterosaurian pycnofibers, though Lingham-Soliar conveniently cuts out the pterosaur part from his quote.&nbsp; Perhaps the greater similarity to pycnofibers and certainty of a Middle Triassic (if not earlier) basal ornithodiran wasn't as insultable as dragging feathers rootward to hypothetical Middle Triassic dinosaurs based solely on quills in one ornithischian?<br /><br />Notably, Lingham-Soliar doesn't seem to believe in pycnofibers either.&nbsp; At least he finally realizes the vast majority, if not all, of pterosaur researchers accept fuzzy pterosaurs (unlike his 2003 paper, or Feduccia from 1996 onward*), though in the end he punts the question- "Given the dismal accounts in the literature of so-called protofeathers in non-avian dinosaurs, e.g., <i>Sinosauropteryx </i>(see Chap. 6), the subject of pycnofibers and their alleged functions such as thermo-regulation and warm-bloodedness will not be discussed any further here."<br /><br />* The funny thing here is he says "Although it was thought to have been based on an unfortunate interpretation of Sharov’s (1971) description of hair-like structures in the pterosaur <i>Sordes pilosus</i>", in reference to the idea of fuzzy pterosaurs, he leaves out who thought that.&nbsp; The answer is himself and Feduccia.&nbsp; But instead of taking responsibility and admitting the BANDits and himself were so grossly ignorant of the pterosaur literature that they misunderstood fuzzy pterosaurs as being based on a misunderstanding of a 1971 paper, he uses the passive "it was thought."&nbsp; "Mistakes were made..."<br /><br /><b>I'm being oppressed!</b> (these titles write themselves) <br /><br />Anyway, on to chapter 7- "The Last Best Hope."&nbsp; Lingham-Soliar actually begins by retelling the story of Galileo's persecution by the Catholic church and of Nazis excluding Jews from scientific research in 1930s Germany.&nbsp; Because Larry Martin died while under house arrest for his views and Alan Feduccia was kicked out of his position at the University of North Carolina, amirite?<br /><br />A section on peer review follows, bemoaning the bias that can exist in the system.&nbsp; The ironic thing here is that I completely agree, but what <b>I</b> find disturbing are all of the falsehoods that slip through peer review in Feduccia's papers.&nbsp; And where are many of those published?&nbsp; In The Auk, published by the American Ornithologist' Union, where Feduccia is a fellow.&nbsp; And who is credited with reviewing e.g. Feduccia's (2013) "Bird origins anew" paper in that journal?&nbsp; Burnham, Czerkas, James, Ruben, Lingham-Soliar, Zhou, and two I haven't heard of.&nbsp; What an unbiased lot!&nbsp; There's some phrase about pots and kettles that seems appropriate here...<br /><br />Page 302 is where things start to get silly- the first figure in this chapter is a cartoon of The Emporer's New Clothes.&nbsp; Because Prum's (2003) statement "current critics of the theropod origin of birds are not doing science" is analogous to telling Feduccia he needs to see a<span style="font-family: inherit;"> nonexistent item, or something.&nbsp; Several figures follow using redrawn scenes from Tim Burton's Alice in Wonderland movie, with quotes changed to deride cladistics.&nbsp; Well if that's not convincing, I don't know what is.&nbsp; Lingham-Soliar then scolds Nature senior editor Henry Gee for a post on the vrtpaleo mailing list where he a</span>ccurately criticized BAND for not using cladistics, correctly stated to be the accepted form of phylogenetic inference in the scientific community.&nbsp; Little did I realize this is "pure censorship."&nbsp; What's especially ironic is Lingham-Soliar's de<span style="font-family: inherit;">scription of this as an inappropriate "outburst" with "purple prose", when Gee wrote that post in reply to Storrs Olson.&nbsp; Has he ever read Olson's writing?!&nbsp; I can't think of anyone in paleontology who writes less professional pieces and gets them published- cladistics was "elevated to a religion years ago", one has to "to suppress one's gag reflex" to read about living dinosaurs, the whole story of Chinese feathered dinosaurs "is essentially a hoax" and "the information has been suppressed" regarding Jehol non-theropods with filaments.&nbsp;&nbsp;</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">But Lingham-Soliar is not to be outdone by Olson.&nbsp; Having already implicitly compared the situation to Galileo and Nazi Germany, he now writes BADists "then state that most scientists support their view, which ironically dominates "primetime" magazines. The regime in apartheid South Africa also said that most people in the country supported its ideology - the fact that most opponents were prevented a voice seemed a tiny, insignificant detail not worth considering."&nbsp; Does he really think there's some population of theropod and basal bird workers out there who believe in BAND but are being prevented from publishing that opinion?&nbsp; All of the theropod workers I know must be very good actors.&nbsp; He even follows Olson's penchant for referring to BAD in religious terms- "people like Dr. Padian with their reduction of science to evangelical preaching and biblical-like hell and damnation for nonbelievers."&nbsp; Who knew BANDits get thrown into an eternal lake of fire?</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">In addition to the Peters-esque conspiracy vibe, Lingham-Soliar is similar to David in claiming his own studies as definitive evidence against Big Paleontology.&nbsp; The following would fit right in on Pterosaur Heresies- "can there be at least some level of justification for Gee’s (2010) euphoric editorial, i.e., if the finding of melanosomes is correct then the filaments are indeed protofeathers? Alas, it seems not as we discovered in Chap. 6 - the notion of melanosomes in the filaments of <i>Sinosauropteryx </i>was shown to be "without scientific merit" (Lingham-Soliar 2011)."&nbsp; Ah yes, 'we' discovered it was shown to be wrong- by me!&nbsp; Also amusing is Lingham-Soliar's constant use of positive adjectives to describe those whose view he supports.&nbsp; Olson is "Curator of Birds (now Emeritus Curator) at The National Museum of Natural History and one of the most eminent ornithologists in the world", while Norell is "curator of dinosaurs at the AMNH."&nbsp; A bit better than a janitor, really.&nbsp; The capitalization, the lack of abbreviation, it's just too good. Similarly, Lingham-Soliar has a penchant for describing his own work with superlatives- "detailed", "rigorous ... which received global attention", "the most revolutionary and explicit images of fiber microstructure in the feather."</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;"><b>BADistics</b></span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Lingham-Soliar writes at length about how cladistics is supposedly unfalsifiable, but that's simply untrue.&nbsp; Any given phylogenetic hypothesis found via cladistic analysis can be falsified by the discovery of a different relationship once new taxa or characters are added.&nbsp; Bullatosauria was popular for a while, the joining of ornithomimosaurs and troodontids recovered in analyses like Holtz (1994).&nbsp; What falsified that?&nbsp; The cladistic analyses of a newly discovered series of basal troodontids (e.g. <i>Sinovenator</i>, <i>Mei</i>) that were more <i>Archaeopteryx</i>-like, not more ornithomimosaur-like.&nbsp; Indeed, we now have taxa like <i>Anchiornis </i>that blur the line so much that some analyses recover th<span style="font-family: inherit;">e</span>m as troodontids and others as birds.&nbsp; The last time I checked the parsimony of alternative hypotheses in the Lori matrix, Bullatosauria was actually the least parsimonious of any previously suggested maniraptoriform topology I could think of.&nbsp;&nbsp;</span><br /><br /><span style="font-family: inherit;">You want falsifiability?&nbsp; If Lingham-Soliar is correct that the filaments of non-bird ornithodirans are collagen (and I agree in a minority of cases they are), then where are all of the fuzzy Jehol fish, frogs, lizards, choristoderes, etc.?&nbsp; These all had collagen in life, so at least a few should be preserved this way if coelurosaurs, pterosaurs and small ornithischians are.&nbsp; No doubt they've been suppressed as Olson claimed. :|&nbsp; Or alternatively, present a repeatable method for distinguishing Jehol feathers on birds/maniraptorans from collagen on other ornithodirans.&nbsp; As far as I know, neither Lingham-Soliar nor anyone else has ever tried to demonstrate Jehol bird filaments are collagen and BANDits have always interpreted them as feathers.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">To continue the hyperbole, did you know that because Ernst Mayr drew attention to pitfalls in cladistics, "he was defamed by supporters of cladistics and the dinosaurian origin of birds in a manner reminiscent of Lysenkoism in the Soviet Union."&nbsp; I must have been too young to remember Mayr being imprisoned in a labor camp and executed.&nbsp; Similarly, "Erik Jarvik had been the butt of the attack by the cladists in the search for tetrapod ancestors" because he had lungfish closer to tetrapods than coelacanths.&nbsp; "It turned out Jarvik was subsequently shown to have been right and the cladists at the time, who were highly critical of him, wrong."&nbsp; So to this day, cladistic analyses of sarcopterygian relationships must be untrusted, right?&nbsp; No, actually what changed the consensus were better cladistic analyses.&nbsp; The more extensive cladistic analyses <u>falsified</u> the hypothesis found in the earlier analyses.&nbsp; Astounding.&nbsp; And you know what's further tested those analyses?&nbsp; Cladistic molecular analyses, which used a completely different character set.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">In case you're wondering, "the Achilles heel of cladistics" was discovered by Jarvik- "cladists arbitrarily pick out unreliable characters from a list without checking their reliability."&nbsp; Of course!&nbsp; Why don't I fire up my objective reliabilitometer more often?!&nbsp; Or if only there were some method to delete and replace random characters then rerun these permutations multiple times, in order to test the strength of each node if certain characters are deleted and other characters are weighted more strongly.&nbsp; But that would be as impossible as pulling yourself up by your own bootstraps!</span><br /><br /><span style="font-family: inherit;"><b>The only good analysis is a BAND analysis </b></span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Repeating the ridiculous notion of Feduccia et al. (2005)- "To take one powerful alleged line of support for the theropod origin of birds namely <i>Caudipteryx</i>, Maryanska et al. (2002) in an exhaustive cladistic analysis showed that rather than a theropod dinosaur as proposed by some cladistics studies, <i>Caudipteryx </i>is a flightless bird. This finding remains undisputed."&nbsp; Undisputed!&nbsp; Except that Lingham-Soliar is blatantly lying, since Maryanska et al. placed <i>Caudipteryx </i>within Theropoda, even using undisputed(!) theropods such as <i>Coelophysis </i>and <i>Allosaurus</i>.&nbsp; And except that the result of placing <i>Caudipteryx</i> closer to Aves than <i>Archaeopteryx </i><u>was</u> disputed by Rauhut (2003), Holtz (2004), Ji et al. (2005), Choiniere et al. (2010) and its variations, Turner et al. (2012), Cau's analysis in Godefroit et al. (2013), Foth et al. (2014), all of the 60+ variants of the TWG matrix published over the last decade and a half, and even James and Pourtless' (2009) terrible BAND analysis.&nbsp; Indeed, I don't think any other analysis has ever recovered that result.&nbsp; As for "exhaustive", Maryanska et al. had 20 taxa and 195 characters, so 3,900 total codings.&nbsp; The analysis of Godefroit et al. had a similar purview but used 101 taxa and 992 characters, resulting in 100,192 codings.&nbsp; Over 25 times the size, and just to make Jarvik happy, it was reweighted two different ways to emphasize those "reliable" characters.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Needless to say, Lingham-Soliar's discussion of James and Pourtless' analysis is completely uncritical.&nbsp; I just <a href="http://theropoddatabase.blogspot.com/2015/01/bandit-cladogram-evaluated-james-and.html">went over why that analysis was so bad</a>, so no need to repeat it here.&nbsp; It's ironic that he complains so often about BAD being 'verificationist' yet just accepts that James and Pourtless had valid, meaningful results despite claiming cladistics is "a discipline I am familiar with as a "working scientist" i.e. when I was involved in taxonomy."&nbsp; What about that <span style="font-family: inherit;">pseudosuchian</span> nested sister to ornithomimosaurs?&nbsp; You didn't notice that because the authors chose not to mention or figure it and you never examined the actual data yourself?&nbsp; I see, I see...</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Lingham-Soliar then complains about a rejection he got, where the reviewer wrote "The author of this paper might be taken more seriously if he did not show such contempt for all other work done on the subject, if he did not ignore all other lines of evidence that he pretends his particular investigation is overthrowing, and if he did not appear so injured by the refusal of the scientific community to recognize his genius."&nbsp; Haha, that describes Theagarten to a tee.&nbsp; "That this was a personal assault did not concern the referee and shamefully was wholly supported by the editor-in-chief of a notable US biological/science journal, with the words, "referees are only human." So was Jack the Ripper!"&nbsp; Why, rejecting poor Lingham-Soliar's manuscript was just like serial murder<span style="font-family: inherit;">!</span>&nbsp; Nazis, Stalin, the Inquisition... what evil characters are left for Lingham-Soliar to equate BADists to?&nbsp; I got it!&nbsp; We're the Galactic Empire and Padian is just like Emperor Palpatine, crushing rebellion and silencing dissenters with his force lightning of parsimony!</span><br /><br /><span style="font-family: inherit;"><span style="font-size: small;"><b>BranchingGate... no wait, Branchgazi</b></span></span><br /><br /><span style="font-family: inherit;"><span style="font-family: inherit;">Alas, <span style="font-family: inherit;">that guess was incorrect</span></span>.&nbsp; Instead, Xu, Prum and Zhao get compared to Richard Nixon and accused of fraud in the next section.&nbsp; Why fraud, you ask?&nbsp; Because Lingham-Soliar doesn't understand how citations work.&nbsp; In the description of <i>Sinornithosaurus</i>' feathers (Xu et al., 2001), they state "the shorter, unbranched integumental appendages of <i>Sinosauropteryx</i>^2, a basal coelurosaur, are also congruent with the predicted Stage I feather morphology."&nbsp; Lingham-Soliar thinks "the superscripted 2 refers to Chen et al. (1998) who they cite as making that critical description namely that feather morphology in <i>Sinosauropteryx </i>are "UNBRANCHED" [my emphasis] as support for their model."&nbsp; And because one line of Chen et al.'s discussion calls the feathers "multibranched", and the last line in the paper suggests feathers evolved from "simpler, branched structures", the authors are "falsifying (OED, misrepresent, distort, (a fact, etc.)) another author’s work."&nbsp; Here's an innocent explanation- that superscript 2 just refers to citing <i>Sinosauropteryx</i>'s latest description, not to any particular interpretation in that description.&nbsp; Why describe its feathers as unbranched?&nbsp; Because Prum (1999) in the paper establishing that model wrote "From my direct observations of the two specimens of <i>Sinosauropteryx </i>(Chen et al., ’98), the integumentary structures appear to consist of unbranched filaments about 20 mm long."&nbsp; Here again, Chen et al. is merely cited as the description, and we can see Prum has his own interpretation of the morphology.&nbsp; He then says "Additional<br />examination of the integumental structures of <i>Sinosauropteryx </i>and <i>Beipiaosaurus </i>is required to establish: (1) whether the filaments are branched, unbranched, or hollow."&nbsp; The truth is that there's still ambiguity, and even the eventual osteology (Currie and Chen, 2001) merely concluded that several features "suggest a feather-like structure with central shafts and plumulaceous barbs" and that "the evidence does favour the interpretation that each has a simple branching structure", but that there was no direct observation of such.&nbsp; Notable is that Lingham-Soliar later quotes this paper in a way I would describe as misleading as it removes the caveats of uncertainty- "They stated that the integumentary structures comprise "central shafts and plumulaceous barbs" and have a "simple branching structure."&nbsp; Who's lying now?</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">It just gets worse from here based on that one misunderstanding of what Xu et al. were citing Chen et al. for in 2001.&nbsp; Xu et al. (2009) claimed <i>Beipiaosaurus</i>' EBFFs were the first unambiguous unbranched feather, yet they "quite astonishingly" cited the 2001 paper as evidence for prior examples being branched!&nbsp; Yeah, because the 2001 paper was primarily describing branched feathers of <i>Sinornithosaurus</i>.&nbsp; The 2001 noting of unbranched feathers in <i>Sinosauropteryx </i>was likely from Prum's 1999 observation, but by 2009 that had been overridden by Currie and Chen's 2001 osteology of the genus, which is indeed cited by Xu et al. in 2009 too.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">But wait!&nbsp; In the slightly later Zheng et al. (2009) description of <i>Tianyulong</i>, "Xing Xu along with his present PNAS co-author Xiaoting Zheng" wrote "In both <i>Tianyulong </i>and <i>Sinosauropteryx</i>, the filamentous structures are singular and unbranched."&nbsp; Ha!&nbsp; They can't keep their story straight!&nbsp; Well, no again.&nbsp; You see, Theagarten, coauthors sometimes disagree about details and even conclusions, and often have different responsibilities.&nbsp; If we check the contributions of each paper, Xu wrote the <i>Beipiaosaurus </i>manuscript, and You wrote the <i>Tianyulong </i>one.&nbsp; Maybe You thinks <i>Sinosauropteryx</i>'s feathers are unbranched and thus disagrees with Currie and Chen but agrees with Prum?</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">So this whole thing is due to Lingham-Soliar not knowing how scientific papers work.&nbsp; It's not fraud.&nbsp; There's no massive coverup.&nbsp; How surprising!</span><br /><br /><span style="font-family: inherit;"><b>Beating a dead <i>Yanornis</i>-<i>Microraptor </i>chimaera </b></span><br /><br /><span style="font-family: inherit;">The next section is a tired recounting of the "Archaeoraptor" story, which Lingham-Soliar should actually be happy about.&nbsp; Nature and Science both rejected manuscripts of the would-be official "Archaeoraptor" description, and the name and chimaerical misinterpretation only made it into the popular press.&nbsp; He writes "Archaeoraptor" "has joined the ranks<br />of paleontological folklore notoriety on a par with Piltdown man", but <i>Eoanthropus </i>(Piltdown man) was accepted by some of the literature for years.&nbsp; Lingham-Soliar further simply believes Olson's statements that Sloan "decided first, that it was appropriate for a journalist to differentiate a taxon new to science and second that evidence claiming to support the new taxon could be presented in a nonpeer reviewed magazine" and that "the name Archaeoraptor liaoningensis Sloan is now<br />available for purposes of zoological nomenclature."&nbsp; Neither is true, as Creisler showed in 2001 on the DML.&nbsp; If you're thinking citing a mailing list would be problematic, Lingham-Soliar is referencing in part a post by Olson on vrtpaleo here, so DML posts would be up for grabs too.&nbsp; Next time check The Theropod Database, Theagarten!&nbsp; I covered the whole story there, with references.&nbsp; Yet due to his misplaced trust in Olson's accuracy, Lingham-Soliar says "one might think it would have spelt the end of Sloan’s days as an editor of NGM."&nbsp; Ouch.</span><br /><b><span style="font-family: inherit;"><br /></span></b><b><span style="font-family: inherit;">Apples to... um... microapples</span></b><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Believe it or not, in this supposedly technical treatise of vertebrate integument, the penultimate section is "Education and Freedom in Apartheid South Africa."&nbsp; That's what it's about too.&nbsp; No paleontology or biology here.&nbsp; Might I humbly suggest that any comparison between BAD vs. BAND and Apartheid is terribly disrespectful to the victims of the latter?</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Theagarten ends the book with a chapter on freedom of press/speech and an appendix which lists the Universal Declaration of Human Rights.&nbsp; If you want yet more irony, recall what I wrote above about the reviewers of Feduccia (2013).&nbsp; Yet Lingham-Soliar has the gall to write "All the evidence points to the fact that in Birds are Dinosaurs only potentially favorable referees are chosen."&nbsp; There's that pot and kettle again.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Actually, it turns out that's not quite how he ends this book.&nbsp; You see, I didn't read ahead when writing this post.&nbsp; I've been writing responses as I read the sections in Lingham-Soliar's book.&nbsp; Remember above, when I guessed "</span><span style="font-family: inherit;">what evil characters are left for Lingham-Soliar to equate BADists to?"&nbsp; My first thought of a remaining evil was Islamic militants.&nbsp; But that was too contemporary for me to feel comfortable joking about, so I went with Star Wars.&nbsp; Yet Lingham-Soliar actually goes there with his Note in Press- "The tragic events in France concerning Charlie Hebdo and Press Freedom is a reminder of those who put their lives on the line for the rights we enjoy in democratic and secular societies - in the words of the children of Soweto [part of his Apartheid section],"the struggle continues." France’s stance against being blackmailed by totalitarian states and terrorists must be applauded lest we become like them."</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Wow.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;"><b>Conclusion</b></span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">The sad part here is that Lingham-Soliar actually has a very good point about how crappy many peer reviews are.&nbsp; You don't have to look further than this blog to find examples of miscoding, not coding, poor procedure, etc..&nbsp; It's in desperate need of revision.&nbsp; But this is hardly unique to BAD.&nbsp; Just look at my recent posts- James and Pourtless' paper is sympathetic to BAND and has terrible flaws, Xu et al.'s 2015 defense of a dromaeosaurid species' validity was conceptually flawed and didn't involve birds, Carrano et al.'s ICZN petition seems shady but is about an <u>actual</u> basal theropod with no bearing on bird origins, Norman's 2014 paper with poor understanding and utilization of phylogenetic nomenclature was about ornithischians, etc..&nbsp; It's not even unique to dinosaurs.&nbsp; In a manuscript I wrote with David Marjanovic criticizing papers that don't code taxa for characters that are known for those taxa, I wrote about dinosaurian examples, and he wrote about basal tetrapod examples.&nbsp; I would bet the issue pervades all of science.&nbsp; Peer reviewers are unpaid<span style="font-family: inherit;">, </span>busy themselves, and often not specialists in what they're reviewing.&nbsp; So they don't check some important things, often don't know what to check for, and are probably often too trusting the author's 'done the work.'&nbsp; It's a problem.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">But what an unhelpful way to present this problem!&nbsp; It proposes a bias in the rigor of peer review, but any BAD supporter who has read a paper by a <span style="font-family: inherit;">BANDit would laugh.&nbsp; Even ignoring any methodological problems with BANDits, they simply don't know much about dinosaurs.&nbsp; As a brief example, here are a couple statements made by Feduccia et al. (2005)- which had Lingham-Soliar as a coauthor.&nbsp; </span>"To illustrate the difficulty of defining the various dinosaur groups, Carroll (1988, p. 290) pointed out that "The 'carnosaur' families may each have evolved separately from different groups that have been classified as coelurosaurs."" "Like the term "thecodont," a collective term to describe Triassic basal archosaurs, coelurosaur and carnosaur describe, respectively, small and large theropod dinosaurs."&nbsp; Did you realize, Theagarten, that these concepts are so obsolete that no one in the 90s, let alone the 2000s, would take them seriously?&nbsp; Carroll's cited work is a general vertebrate paleontology textbook written 17(!) years earlier.&nbsp; Who would ever think that's a valid source for contemporary taxonomy?!&nbsp; Past the 80s, Coelurosauria has basically always been recognized as the clade of theropods closer to birds than to carnosaurs, including <i>Compsognathus</i>, <i>Ornitholestes</i>, ornithomimosaurs, oviraptorosaurs, dromaeosaurids and troodontids.&nbsp; Even if you dispute birds belong there, the concept<span style="font-family: inherit;"> of the group has remained consistent.&nbsp; The idea that carnosaurs are big theropods and coelurosaurs are small theropods is the kind of simplification of 1920-1970s phylogeny that you read in old childrens books.&nbsp; So how did such a blatant misrepresentation make it through peer review into your paper?&nbsp; And there are endless examples like this in BANDit works.&nbsp; I could say that you and Feduccia are maliciously misrepresenting how well established the consensus of dinosaur phylogeny/taxonomy is.&nbsp; The same paper also says "</span>the question of whether birds are derived from dinosaurs depends on what one defines as dinosaur, or the Dinosauromorpha", even though the concept of Dinosauromorpha has <u>never</u> been controversial, nor is contingent on the definition of Dinosauria, which itself hasn't been controversial since dinosaur monophyly was established in the 70s.&nbsp; Is this because the BANDits have peer reviewers that are in cahoots to let bad information through?&nbsp; No.&nbsp; It's just that peer reviews are sometimes poor and that you and Feduccia don't know much about dinosaurs.&nbsp; "Basal dinosaur" <i>Sinosauropteryx</i>... "undisputed" Maryanska et al. 2002...</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Next up is your mischaracterization of Prum's (2003) "not doing science" critique as being focused on cladistics.&nbsp; Far from it.&nbsp; Prum was criticizing Feduccia for- not explicitly proposing an alternative phylogenetic position for birds (e.g. why or why not are they in Avemetatarsalia, or Archosauria, or Archosauriformes); ignoring how character evidence works together to support or refute hypotheses, instead only taking single characters and disputing their homology, stating a single character is homoplasious, and ignoring the fact convergence can only be shown if the majority of characters support the real relationship; and most importantly for completely contradicting all of his previous work which disputed similarity between dromaeosaurids and birds without any sort of explanation for how he got things so wrong before, and why we should trust his methods now that they've been disproven.&nbsp; Was your mischaracterization malicious or merely ignorant?</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Your paper has continual complaints about free speech, free press, censorship!&nbsp; But BANDits get tons of media presence.&nbsp; If we look at everything since Gee's 1999 "censorship" mailing list post, Feduccia's gotten published on the topic of bird origins...</span><br /><span style="font-family: inherit;">- a 2000 American Zoologist article.&nbsp;</span><br /><span style="font-family: inherit;">- a 2001 Journal of Ornithology article.</span><br /><span style="font-family: inherit;">- a 2001 rebuttal in TREE.</span><br /><span style="font-family: inherit;">- a 2002 Auk article.&nbsp;</span><br /><span style="font-family: inherit;">- a 2002 Naturwissenschaften article.</span><br /><span style="font-family: inherit;">- a 2003 TREE letter.</span><br /><span style="font-family: inherit;">- a 2003 Auk book review.</span><br /><span style="font-family: inherit;">- a 2005 Journal of Morphology article.</span><br /><span style="font-family: inherit;">- a 2007 Auk article.</span><br /><span style="font-family: inherit;">- a 2007 Proceedings B article.</span><br /><span style="font-family: inherit;">- a 2013 Auk article.</span><br /><span style="font-family: inherit;">- a 2014 Journal of Ornithology Article.<br />And more, no doubt.&nbsp; There is an article published almost every year.&nbsp; He also wrote the book "Riddle of the Feathered Dragons: Hidden Birds of China" from New Yale University Press.&nbsp; How is Feduccia being prevented from airing his views?&nbsp; That's not an outlier either.&nbsp; Martin got plenty of publications, as did Czerkas.&nbsp; And just look at yourself, Theagarten.&nbsp; You've had numerous articles published disputing BAD interpretations.&nbsp; Some were rejected, but EVERYONE gets many manuscripts rejected.&nbsp; And you got this entire present volume published!&nbsp; You got to air your accusatory views of a conspiracy in science, via a scholarly publisher.&nbsp; How is this being censored?&nbsp; Freedom of speech means you're allowed to say what you want without being a criminal, freedom of press means you're allowed to publish what you want without being a criminal.&nbsp; Neither means any particular platform, e.g. Nature, has to host your views<span style="font-family: inherit;">.</span></span><br /><br /><span style="font-family: inherit;"><span style="font-family: inherit;">You want to <span style="font-family: inherit;">accuse BADists of fraud, but in this very volume you fr<span style="font-family: inherit;">ame the<span style="font-family: inherit;"> <i>Sinosauropteryx </i><span style="font-family: inherit;">media rush in 1996 as if it pr<span style="font-family: inherit;"><span style="font-family: inherit;">e</span>ceded&nbsp; </span></span></span></span></span>its description ("Soon after [in 199<span style="font-family: inherit;">6]</span>,<span style="font-family: inherit;"> </span>without any scientific investigation, a pen-and-ink sketch of the sensational specimen<span style="font-family: inherit;"> </span>appeared on the front page of The New York Times, as support for the theory.<span style="font-family: inherit;">&nbsp; </span>In 1998, a description of <i>Sinosauropteryx </i>appeared in the journal Nature (Chen<span style="font-family: inherit;"> </span>et al. 1998).").&nbsp; You know <i>Sinosauropteryx </i>wa<span style="font-family: inherit;">s <span style="font-family: inherit;">officially described, as feathered, in 1996 by Ji and Ji, right?&nbsp; You f<span style="font-family: inherit;">alsely state Ma<span style="font-family: inherit;">ry<span style="font-family: inherit;">anska et al. (2002) did no<span style="font-family: inherit;">t recover <i>Cau</i><span style="font-family: inherit;"><i>dipteryx </i>as a theropod, and that its results are undisputed more than a decade later.&nbsp; You conve<span style="font-family: inherit;">nien<span style="font-family: inherit;">tly cut out reference to pterosa<span style="font-family: inherit;">urs in Xu et al.'s (2009) quote.&nbsp; <span style="font-family: inherit;">You repeat Olson's incorrect allegations "Archaeoraptor" was <span style="font-family: inherit;">intended to be established by Sloan in <span style="font-family: inherit;">National Geog<span style="font-family: inherit;">raphic and that his article was valid under the ICZN.&nbsp; Should I accuse you of fraud, or are you just <span style="font-family: inherit;">very ignorant of dino<span style="font-family: inherit;">saur res<span style="font-family: inherit;">earch, and never bothered to look up <i>Sinosauropte</i><span style="font-family: inherit;"><i>ry</i><span style="font-family: inherit;"><i>x</i>'s Chinese description, don't follow <span style="font-family: inherit;">theropod phylogenetics papers, didn't bother researching "Ar<span style="font-family: inherit;">chaeoraptor" past what Olson said,<span style="font-family: inherit;"> etc.?</span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span></span> </span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">Finally, it turns out I was wrong.&nbsp; I wrote in reference to Olson that "</span><span style="font-family: inherit;">I can't think of anyone in paleontology who writes less professional pieces and gets them published."&nbsp; That was before I read the rest of this document.&nbsp; Feduccia and Martin display gross ignorance of dinosaur knowledge, Olson has a sharp tongue and tedious fondness for religious analogies, but it's only you, Theagarten, who has compared BAD researchers to the Inquisition, Nazis, Apartheid, Stalin and Al-Qaeda.&nbsp; That's beyond the pale and completely unprofessional.&nbsp; Even if everything you wrote were true and there was some massive conspiracy to only allow BAD ideas into Nature/Science/PNAS and the popular media, that would be insignificant compared to the evil of those parties.&nbsp; Shame on you.&nbsp; And a hearty **** you on behalf of Xu, Prum and Zhao who you directly accuse of fraud.&nbsp; Grow up and get serious.</span><br /><span style="font-family: inherit;"><br /></span><span style="font-family: inherit;"><b>Reference</b>- Lingham-Soliar, 2015. The Vertebrate Integument<span style="font-family: inherit;"> </span>Volume 2<span style="font-family: inherit;">: </span>Structure, Design and Function. Springer<span style="font-family: inherit;">,</span> Heidelberg New York Dordrecht London. 348 pp.</span><br /><br />Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com18tag:blogger.com,1999:blog-3248412803814730250.post-64832587035668144782015-01-24T04:30:00.000-08:002015-01-25T01:49:04.714-08:00BANDit cladogram evaluated - James and Pourtless 2009Besides Hou et al. (1996), the only BANDit (Birds Are Not Dinosaurs) cladistic analysis has been that of James and Pourtless (2009).&nbsp; Little has been said about this paper, with most of the commentary noting the backwards philosophy the authors had in coding characters with supposedly controversial homology as unknown instead of letting phylogenetic analysis establish homology.&nbsp; Well I finally looked at it in a little detail, and found a far larger issue.<br /><br />First, let's go over the basic tests and conclusions of James and Pourtless' study.&nbsp; After making the almost correct observation that no published cladistic analysis has tested whether birds are dinosaurs (Senter 2004 did, but while the authors cite it they don't comment on it in that regard), James and Pourtless aim to correct this by constructing their own dataset with alternative proposed bird ancestors included.&nbsp; With basal archosauriforms as the outgroup, the authors also include crocodylomorphs,&nbsp; <i>Longisquama,</i> a few other pseudosuchians and numerous theropods.&nbsp; Running their dataset of 242 characters and 79 taxa, they recover a large number of MPTs.&nbsp; These are only ever illustrated as majority rule bootstrap trees after a posteriori pruning of taxa, and each alternative tree is basically a well supported Archosauria with a massive polytomy between crocodylomorphs, numerous theropod lineages and birds.&nbsp; <i>Longisquama </i>weakly (55% bootstrap) groups with birds if 21 "controversially homologous" characters (which are only coded for non-dinosaurs) are excluded and non-bird maniraptorans are pruned away.&nbsp; Statistical tests were done, seemingly showing the 'basal archosaur' (= <i>Longisquama</i>), crocodylomorph and dinosaur hypotheses of bird origins aren't any worse than each other.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-s5Wl7l-VQA4/VMNJnzWn_nI/AAAAAAAAAlE/0wjfJYmbrWo/s1600/James%2BPourtless%2Bclado.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" src="http://4.bp.blogspot.com/-s5Wl7l-VQA4/VMNJnzWn_nI/AAAAAAAAAlE/0wjfJYmbrWo/s1600/James%2BPourtless%2Bclado.jpg" height="243" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Figure 12 of James and Pourtless (2009), showing majority rule bootstrap tree with numerous taxa including all maniraptorans pruned away a posteriori.&nbsp; *Gasp* It looks like birds are closer to non-dinosaur <i>Longisquama </i>and that even crocs could be as close to birds as dinosaurs.&nbsp; Were BANDits right all along? (Spoiler- NO)</td></tr></tbody></table>How to explain this?&nbsp; First, I should note that when you run their data, the resulting MPTs are pretty standard.&nbsp; As shown in my typed cladogram below, while there are plenty of weird details going on in Theropoda, the basic structure of birds within Pennaraptora within Maniraptora within Coelurosauria within Theropoda within Saurischia within Archosauria is present.&nbsp; The only odd relevant aspects are <i>Effigia </i>is an ornithomimosaur and <i>Longisquama </i>is deeply embedded in Pennaraptora.&nbsp; Note there's no way of knowing <i>Effigia </i>groups with ornithomimosaurs by reading the paper, as its exact results aren't mentioned and it's always pruned out of the figured trees.&nbsp; I find this suspicious.&nbsp; In any case, <i>Longisquama </i>being sister to ornithines is discussed since this is the current favored alternative of BANDits.&nbsp; Besides the paltry literature, the authors depended on photos at the KU for their coding of the genus, and present a cranial reconstruction.&nbsp; This reconstruction looks as if it was done assuming a theropodan morphology for the taxon.&nbsp; For example, there's a T-shaped structure at the anterodorsal corner of the orbit, interpreted as a lacrimal by the authors.&nbsp; But it would also compare well to a prefrontal in a basal diapsid like <i>Coelurosauravus</i>, where the lacrimal is reduced.&nbsp; Similarly, the strip under the supposed supraorbital ridge (yellow) would match the posterodorsal strip of the lacrimal/prefrontal in <i>Coelurosauavus</i>.&nbsp; Or the wedge just behind this, while unidentified by the authors, would match the postfrontal of <i>Coelurosauravus</i>.&nbsp; The supposed antorbital fossa (red) matches the external naris of <i>Coelurosauravus </i>well, and even the big posterodorsal hole ("?" in their figure) would match <i>Coelurosauravus</i>' if it's a supratemporal fenestra as identified by most previous authors.&nbsp; This isn't to say that <i>Longisquama</i> is a close relative of <i>Coelurosauravus </i>(as Senter 2004 recovered), merely that the lens you view a fossil like <i>Longisquama</i> through based on photos can affect your identifications.&nbsp; The controversial aspects of <i>Longisquama</i>'s anatomy are similarly all coded as theropod-like- thecodont teeth, antorbital fenestra with maxillary fenestra, furcula, etc..<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://2.bp.blogspot.com/-wIpLjzT9syA/VMNZPX3oStI/AAAAAAAAAlY/RgZd0y40OfE/s1600/Longisquama%2Bskull%2BJames%2Band%2BPourtless%2B2009%2Bvs%2BCoelurosauravus.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" src="http://2.bp.blogspot.com/-wIpLjzT9syA/VMNZPX3oStI/AAAAAAAAAlY/RgZd0y40OfE/s1600/Longisquama%2Bskull%2BJames%2Band%2BPourtless%2B2009%2Bvs%2BCoelurosauravus.jpg" height="221" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Longisquama </i>skull (counterslab of PIN 2584/4) and interpretation of James and Pourtless (2009) on top.&nbsp; <i>Coelurosauravus </i>skull reconstruction at bottom after Evans and Haubold (1987), modified so that the entire spiked temporal element is the squamosal after Schaumberg et al. (2007).&nbsp; I see the supposed antorbital fossa (afo in red), but don't think its obviously a fossa, as opposed to a fenestra (e.g. the naris) or bone process itself.&nbsp; Similarly, the entire ventral edge of the supposed antorbital fenestra (aof) looks like a crack to me, including the supposed dorsal process of the maxilla (dpm), supposedly there to support an interfenestral bar.&nbsp; I don't see any posterior edge of the antorbital fenestra either, so that that whole area could be a solid maxilla.&nbsp; The ventral process (vp) is apparent, but if anything seems separated from the nasal by a suture (shown by the green space).</td></tr></tbody></table><div class="separator" style="clear: both; text-align: center;"></div>As for the matrix, there are of course lots of miscodings (e.g. character 5 is for dermal armor, which is coded unknown for almost all theropods and <i>Effigia</i>, plus coded absent in <i>Ceratosaurus</i>, the one theropod that actually has it).&nbsp; Some of the characters are formed terribly while others are partitioned in a way that weights them (e.g. each dentigerous element has its own character for tooth serrations, each a three state ordered character of the form '0- all serrated; 1- some serrated; 2- none serrated'; thus any taxon leading from serrated to unserrated teeth needs six steps to do so).&nbsp; The 21 "controversially homologous" characters include every manual character due to the I-II-III vs. II-III-IV issue, though oddly these aren't even coded for dinosaurs with five digits like <i>Herrerasaurus</i>.&nbsp; There's also a huge imbalance of taxa, with a whopping 45 non-avian theropods plus 16 birds, only 9 pseudosuchians, 3 non-archosaurs, <i>Eoraptor</i>, <i>Marasuchus </i>and <i>Scleromochlus</i>.&nbsp; These are all issues, but not the most important one.<br /><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">|--Proterosuchus<br />`--+--Euparkeria<br />...`--+--Erythrosuchus<br />......`--+--Pseudosuchia(Ornithosuchus,Postosuchus,Crocodylotarsi)<br />.........|--Scleromochlus<br />.........|--Marasuchus<br />.........`--+--Eoraptor<br />............`--+--Herrerasaurus<br />...............|--Carnosauria("Syntarsus",Dilopho,Cerato,Sinraptor,Allo,Tyranno)<br />...............`--+--Juravenator<br />..................|--Sinosauropteryx<br />..................|--Compsognathus+Huaxiagnathus<br />..................|--Guanlong+Dilong<br />..................|--<b>Effigia</b>+Ornithomimosauria<br />..................|--Microvenator<br />..................`--+--Coelurus<br />.....................|--Ornitholestes<br />.....................|--Falcarius<br />.....................`--+--Therizinosauroidea<br />........................`--+--Oviraptorosauria+Dromaeosauridae<br />...........................`--+--Troodontidae<br />..............................`--+--Pelecanimimus+Avimimus+Parvicursorinae<br />.................................`--+--Caudipteryx<br />....................................`--+--<b>Longisquama</b><br />.......................................`--Ornithes </span><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;"></span><br /><div style="text-align: center;"><span style="font-size: x-small;">Actual strict consensus of James and Pourtless' data, simplified so that genera which form monophyletic clades are represented by their clade names.&nbsp; Note no dinosaurs were coded for manual characters, <i>Effigia </i>is an ornithomimosaur, and <i>Longisquama </i>is deeply nested in Theropoda.</span></div><br />No, the biggest problem with James and Pourtless' analysis is that its matrix consists mostly of characters designed to diagnose coelurosaur clades (e.g. 96 from Clark et al.'s 2002 TWG analysis; 26 from Chiappe's 2002 bird analysis) plus those suggested by BANDits to group <i>Longisquama </i>and crocs with birds.&nbsp; So what happens when you analyze <i>Longisquama</i> in a coelurosaur/bird matrix after reconstructing its skull with theropod presumptions and pretending no dinosaur preserves hands?&nbsp; Given it doesn't preserve sacrum, pelvis, hindlimbs or tail, and that the vertebrae are basically uncodable, it emerges as a coelurosaur.&nbsp; When chatting with Nick Gardner about this I joked "I bet if you coded e.g. <i>Coelurosauravus</i>' front half into the matrix, it would be coelurosaurian too."&nbsp; So I coded <i>Coelurosauravus</i>' front half, the same parts preserved for <i>Longisquama</i>, and lo...<br /><br /><span style="font-family: &quot;Courier New&quot;,Courier,monospace;">|--Proterosuchus<br />`--+--Euparkeria<br />...`--+--Erythrosuchus<br />......`--+--Pseudosuchia(Sclero,Ornithosu,Posto,Crocodylo)<br />.........`--+--Marasuchus<br />............`--+--Eoraptor<br />...............`--+--Herrerasaurus<br />..................`--+--Carnosauria("Syntarsus",Dilopho,Cerato,Sin,Allo,Tyranno)<br />.....................`--+--Compsognathidae<br />........................`--+--Guanlong+Dilong<br />...........................`--+--Microvenator<br />..............................`--+--Ornitholestes<br />.................................`--+--+--<b>Coelurosauravus</b><br />....................................|..`--+--<b>Effigia</b><br />....................................|.....`--Ornithomimo (inc. Pele,<b>Longisquama</b>)<br />....................................`--+--Coelurus<br />.......................................`--+--Falcarius<br />..........................................`--+--Therizinosauroidea<br />.............................................`--+--Ovirapt(inc.Caudi)+Dromaeo<br />................................................`--+--+--Troodontidae<br />...................................................|..`--Avimimus+Parvicursorinae<br />...................................................`--+--Sinovenator<br />......................................................`--Ornithes</span><br /><br />I won my own bet.&nbsp; Interestingly, this tree also has a lot more resolution and details which agree with the consensus (e.g. <i>Pelecanimimus </i>in Ornithomimosauria, <i>Caudipteryx </i>in Oviraptorosauria), and <i>Longisquama </i>is moved from Avialae to Ornithomimosauria.&nbsp; Note <i>Coelurosauravus </i>is not thought by anyone to have anything to do with dinosaurs or birds, but still ends up as a maniraptoriform in James and Pourtless' matrix.&nbsp; This fairly neatly proves my idea that <i>Longisquama </i>could be a far more basal diapsid and still emerge as a coelurosaur.&nbsp; Note too that <i>Longisquama </i>is also an ornithomimosaur once the front half of <i>Coelurosauravus </i>is included, perhaps suggesting more signal between them than between <i>Longisquama </i>and birds.<br /><br />So my conclusion is that very few characters were included that would support e.g. Tetanurae, Avepoda, Theropoda, Saurischia, Dinosauromorpha.&nbsp; Just enough are included to get a basically consensus phylogeny even without manual characters, though not enough to properly place <i>Effigia </i>or the front half of <i>Coelurosauravus</i>.&nbsp; The latter plus the theropodan assumptions in <i>Longisquama</i>'s anatomy makes placement of thus genus particularly untrustworthy.&nbsp; Certainly, not enough characters were included to survive bootstrap analysis, where random characters are deleted or repeated and the analysis is rerun.&nbsp; It doesn't mean that "both the "early-archosaur" and "crocodylomorph" hypotheses are at least as well supported as the BMT [BAD] hypothesis", it means that James and Pourtless made a crappy analysis.<br /><br /><b>References</b>-&nbsp; Evans and Haubold, 1987. A review of the Upper Permian genera <i>Coelurosauravus</i>, <i>Weigeltisaurus </i>and <i>Gracilisaurus </i>(Reptilia: Diapsida). Zoological Journal of the Linnean Society. 90(3), 275-303.<br /><br />Hou, Martin, Zhou and Feduccia, 1996. Early adaptive radiation of birds: Evidence from fossils from Northeastern China. Science. 274, 1164-1167.<br /><br />Senter, 2004. Phylogeny of the Drepanosauridae (Reptilia: Diapsida). Journal of Systematic Palaeontology. 2, 257-268.<br /><br />Schaumberg, Unwin and Brandt, 2007. New information on the anatomy of the Late Permian gliding reptile <i>Coelurosauravus</i>. Palaontologische Zeitschrift. 81(2), 160-173. <br /><br />James and Pourtless, 2009. Cladistics and the origin of birds: A review and two new analyses. Ornithological Monographs. 66, 78 pp.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-64686679146793091062015-01-15T03:27:00.000-08:002015-01-15T03:27:31.763-08:00Is Linheraptor a synonym of Tsaagan?One of the first theropod papers of 2015 is Xu et al.'s defending <i>Linheraptor </i>as being distinct from <i>Tsaagan</i>, after they were synonymized by Senter (2011) and Turner et al. (2012).&nbsp; I provisionally synonymized them on the Database because Turner's arguments seemed sound, though I haven't actually studied the problem myself.&nbsp; One interesting aspect here is that this paper is basically a criticism of the details and methods of Turner et al., which was basically a published version of part of Turner's (2008) thesis.&nbsp; Yet Norell is a coauthor, who was not only a coauthor of Turner et al., but also Turner's advisor for his thesis.&nbsp; <br /><br />Unfortunately, after reading Xu et al.'s arguments, my conclusion is that a LOT of work went into arguing a point no one disputes- the holotypes are not identical.&nbsp; The authors make a huge point of the number of characters they found which differ between the holotypes- sixty-one.&nbsp; Also that many of these are found in some dromaeosaurids, but not others.<br /><br />Xu et al. correctly state "Proposals of synonymy are usually based on a judgment that putative diagnostic differences between the taxa in question are individual, ontogenetic, or sexually dimorphic variations rather than taxonomic ones, making one of the new taxa invalid. Taxonomically informative variations can sometimes be distinguished from intraspecific ones if a sample large enough to provide a basis for rigorous investigation of patterns of variation is available."&nbsp; They then say "In the present case, Evans et al. (2013), Senter et al. (2012) and Turner et al. (2012) attributed the proposed diagnostic features distinguishing <i>L. exquisitus</i> from <i>T. mangas</i> to individual intraspecific variation, but avenues for testing this hypothesis are limited because each species is currently represented only by a single individual."<br /><br />Well, no.&nbsp; You don't need the exact species under consideration to study the extent of individual variation, you can use relatives.&nbsp; Turner himself had the right idea to use the multiple known skulls of the closely related <i>Velociraptor mongoliensis</i> as the basis for judging differences between <i>Tsaagan </i>and <i>Linheraptor</i>.&nbsp; Xu et al.'s response to this is rather comical- "a comprehensive taxonomic review [of <i>Velociraptor</i>] has yet to be published to determine how observed morphological variations relate to inter- or possibly intraspecific factors. This can be addressed by including all <i>Velociraptor </i>specimens in a specimen-level phylogenetic analysis, by using morphometric methods to quantify the variation present and by deepening our understanding of the biological significance of the variations observed. Until this work has been completed it is in our view that noticeable variations between <i>L. exquisitus</i> and <i>T. mangas</i> are grounds for taxonomic separation."<br /><br />So until we perform these studies which have been published so far for zero (specimen-level phylogenetic analysis), ?five (using morphometric methods to quantify the variation present) and ?zero (deepening our understanding of the biological significance of the variations observed) Mesozoic theropods on <i>Velociraptor</i>, we can't treat multiple specimens of it as one taxon?&nbsp; That's a bit of a far goalpost, don't you think?&nbsp; The morphometric studies done on other theropods haven't even resolved their taxonomy (e.g. <i>Allosaurus</i>, <i>Archaeopteryx</i>), so I'm skeptical a similar study on <i>Velociraptor </i>would yield useful results either.&nbsp; If we were to extend Xu et al.'s demands to other taxa, we'd basically have to say we know nothing yet of intraspecific variation in Mesozoic theropods.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://4.bp.blogspot.com/-BgZBjduxymg/VLedGciJVkI/AAAAAAAAAko/7qa4daK3mtw/s1600/Tsaagan%2BLinheraptor%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" src="http://4.bp.blogspot.com/-BgZBjduxymg/VLedGciJVkI/AAAAAAAAAko/7qa4daK3mtw/s1600/Tsaagan%2BLinheraptor%2Bcomp.jpg" height="320" width="227" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Differences between the <i>Tsaagan </i>(first and third rows) and <i>Linheraptor </i>(second and fourth rows) holotypes. After Xu et al. (2015).</td></tr></tbody></table>Xu et al. include extensive comparisons to other dromaeosaurid taxa for each difference noted.&nbsp; They say "Taken together, the distributions of these features among dromaeosaurids not only demonstrate that <i>L. exquisitus</i> is a valid taxon distinct from <i>T. mangas</i> but also provide important information on dromaeosaurid interrelationships."&nbsp; Yet in almost every case, there's only one skull known for each other taxon as well.&nbsp; Thus if e.g. <i>Tsaagan </i>shares a character with the <i>Dromaeosaurus</i> holotype not found in <i>Linheraptor</i>, yes it <u>could</u> mean that's a phylogenetically useful character uniting the first two genera, but it <u>could also</u> mean it's an individually variable character that the one preserved specimen of <i>Dromaeosaurus </i>happens to share with the only specimen of Tsaagan and not the only specimen of <i>Linheraptor</i>.<br /><br />You'd have to see how the character distribution worked out in total, as if e.g. <i>Tsaagan </i>and <i>Dromaeosaurus </i>shared a lot of derived characters not found in <i>Linheraptor</i>, that would be increasingly good evidence for a phylogenetic signal (or sexual dimorphism or ontogenetic change, I suppose) instead of coincidence.&nbsp; Needless to say, Xu et al. do not do this and indeed find even more characters shared between <i>Tsaagan </i>and <i>Linheraptor</i>.<br /><br />My basic response to Xu et al. is "Could I find 61 comparable differences between two skulls and necks of specimens near universally agreed to be <i>Tyrannosaurus rex</i>? Yes I could."&nbsp; Indeed, in every example I've looked into, from <i><a href="http://theropoddatabase.com/Carnosauria.htm#Allosaurusfragilis">Allosaurus</a> </i>to <i><a href="http://theropoddatabase.com/Dromaeosaurs.htm#Microraptorzhaoianus">Microraptor</a> </i>to <a href="http://theropoddatabase.com/Avialae.htm#Archaeopteryxlithographica"><i>Archaeopteryx</i></a>, every specimen differs in numerous other ways from every other.&nbsp; Importantly, the differences never sort themselves into mutually exclusive pools, so that instead of Morph A having traits ABCD and Morph B having traits 1234, we instead get a specimen with 1B34, one with 123D, one with A2C4, one with 12C4, etc..&nbsp; So either every known relatively complete and described Mesozoic theropod individual is its own species, or Mesozoic theropods had significant individual variation, just like living organisms.&nbsp; I know which option I choose.<br /><br /><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody><tr><td style="text-align: center;"><a href="http://3.bp.blogspot.com/-1AmJf0q8ZIY/VLeedOztuVI/AAAAAAAAAk0/_WkvvgYmQYw/s1600/Velociraptor%2Bcomp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" src="http://3.bp.blogspot.com/-1AmJf0q8ZIY/VLeedOztuVI/AAAAAAAAAk0/_WkvvgYmQYw/s1600/Velociraptor%2Bcomp.jpg" height="320" width="261" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Variation in three <i>Velociraptor mongoliensis</i> skulls. Top- holotype AMNH 6515, middle IGM 100/25 (fighting specimen), bottom IGM 100/982.&nbsp; Note the numerous differences (anterior premaxillary angle in holotype, maxillary fenestra shape in top two, anterior antorbital fenestra shape, ventral lacrimal process angle in 982, posterior jugal outline in holotype, etc.).&nbsp; After Norell et al. (2006).</td></tr></tbody></table><br /><br />To finish up the criticisms, I find Xu et al.'s statement that "In some taxonomic studies, some taxonomic indicators have been based on superficial rather than detailed morphological comparisons" to be based on problematic examples.&nbsp; They say "<i>Saurornitholestes </i>and <i>Deinonychus </i>have been regarded as junior synonyms of <i>Velociraptor </i>based on overall similarities (Paul, 1988), although both are now widely accepted as valid taxa (e.g., Turner et al., 2012)."&nbsp; But Paul never advocated those as being the same species, merely different species in one genus.&nbsp; That's a completely different issue, since genera are (more) subjective and Turner's analysis even finds these to be monophyletic to the exclusion of <i>Dromaeosaurus</i>, the only other decently described dromaeosaurid Paul knew of (Paul questioningly includes <i>Adasaurus </i>as a dromaeosaurine but notes "not enough has been published for us to be certain of anything").&nbsp; If the authors wanted an example of bad Paul synonymization, they had the easy target of <a href="http://theropoddatabase.blogspot.com/2010/10/princeton-field-guide-to-dinosaurs_07.html">his 2010 synonymization of <i>Tsaagan </i>with <i>Velociraptor</i></a>, but not <i>Deinonychus </i>or <i>Saurornitholestes</i>.<br /><br />"Similarly, the basal tyrannosauroid <i>Guanlong </i>has been suggested to be a sub-adult individual of the basal tetanuran <i>Monolophosaurus </i>based on the fact that the two taxa are both characterized by a cranial crest and several other superficially similar features (Carr, 2006), though this proposal has received little acceptance (Brusatte et al., 2010, 2012)."&nbsp; Well, Carr's argument was only an SVP abstract, so it hasn't had a chance to be detailed yet.&nbsp; Seems like a low blow.<br /><br />"In addition to considering <i>L. exquisitus</i> to be a synonym of <i>T. mangas</i>, Turner et al. (2012) scored some character states for <i>[Sinornithosaurus] millenii</i> based on IVPP V 169041). However, V 16904 is not referable to <i>S. millenii</i> for two key reasons. First, V 16904 is inferred to be more ontogenetically advanced than the holotype specimen of <i>S. millenii</i> (V 12811) based on fusion features (the neurocentral sutures are fully closed in the preserved vertebrae of V 16904 but are evident in at least some vertebrae of V 12811, the proximal tarsals are fused to the tibia in V 16904 but remain separate in V 12811), but is significantly smaller than the latter specimen; and second, V 16904 differs from the holotype of <i>S. millenii</i> in some important morphological features, such as the fact that all of the teeth lack denticles in the former specimen but have denticles in the latter."&nbsp; Here we have a brief example of the splitter mindset- differences equal taxonomic separation.&nbsp; For just a few counter-examples- the <i>Mei </i>referred specimen has a tibiotarsus and fused presacral neurocentral sutures but is 80% the size of the holotype that doesn't; the <i>Sinovenator </i>paratype has a tibiotarsus but is 89% the size of the holotype that doesn't; <i>Microraptor</i> specimens can have serrations on both carinae (NGMC 00-12-A), only distal serrations (holotype) or no serrations (IVPP V13320).&nbsp; Again, either we have a ton of unrecognized coexisting species, or species exhibit variation.<br /><br />As far as good points go, besides more details and gorgeous photos of the amazingly complete <i>Linheraptor </i>skull (e.g. both scleral rings are articulated and in place), Xu et al. are right to criticize <a href="http://theropoddatabase.blogspot.com/2012/08/turner-et-al-2012-great-review-sloppy.html">some sloppiness in Turner's work</a>.&nbsp; But they missed out on the least justified problem Turner had regarding <i>Linheraptor</i>- if he viewed it as synonymous with <i>Tsaagan</i>, WHY didn't he code the basically complete <i>Linheraptor </i>specimen for <i>Tsaagan</i>'s OTU?!<br /><br />In conclusion, Xu et al. state "... the contrast between our own perception of <i>L. exquisitus</i> as a valid taxon with many distinguishing features and the view that <i>L. exquisitus</i> is a junior synonym of <i>T. mangas</i> (Evans et al., 2013; Senter et al., 2012; Turner et al., 2012) presents an example of the principle that “similarity lies in the eyes of the beholder” (Clark, 1992)."&nbsp; As seven of the eight authors of this paper were authors of the <i>Linheraptor </i>description, I might be forgiven for wondering if "similarity lies in the eyes of the namer" has some relevance as well. :)<br /><br /><b>References</b>- Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp. <br /><br />Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis. Columbia University. 666 pp. <br /><br />Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24, 2197-2216.<br /><br />Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.&nbsp; <br /><br />Xu, Pittman, Sullivan, Choiniere, Tan, Clark, Norell and Wang, 2015. The taxonomic status of the Late Cretaceous dromaeosaurid <i>Linheraptor exquisitus</i> and its implications for dromaeosaurid systematics. Vertebrata PalAsiatica. 53(1), 29-62. Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com0tag:blogger.com,1999:blog-3248412803814730250.post-13758436651294563972015-01-10T03:04:00.005-08:002015-01-10T03:04:34.246-08:00Big "New Years" Theropod Database updateHappy only over a week from New Years everyone!&nbsp; Time for a Theropod Database update.&nbsp; Besides all of the new taxa, there's a huge revising of Morrison allosaurids and Baharija taxa (check out the tetanurine <i>Bahariasaurus</i>).&nbsp; Enjoy all of the <a href="http://theropoddatabase.com/Updates.htm">updates</a>.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com0